Pacific Science (1997), vol. 51, no. 4: 424-439 © 1997 by University of Hawai'i Press. All rights reserved

Dispersal and Vicariance in Hawaiian Platynine Carabid (Coleoptera)1

JAMES K. LIEBHERR2

ABSTRACT: The monophyletic, native Hawaiian have diversified on the Hawaiian Island chain through progressive colonization, mixed with vicariance on the various islands. Single-island endemism stands at 97% of the species, with the few widespread species exhibiting distributions largely congruent with the fundamen­ tal area cladogram found using cladistic biogeographic methods. The cost of accepting an ad hoc dispersal hypothesis for individual taxa that conflicts with the fundamental area cladogram is weighed against the savings in items of error when taxa are excluded from the biogeographic analysis. Based on this objective assess­ ment, only one back-dispersal from Maui Nui to O'ahu is supported. Vicariance of Maui Nui, leading to the present-day islands of Moloka'i, Uina'i, and Maui, has resulted in seven resolvable species triplets composed of single-island endemics occupying these areas. These seven triplets represent five biogeographic patterns, necessitating explanation by numerous ad hoc hypotheses of extinction to support a single hypothesis of area relationships. In six of the seven triplets, the cladistically basal species exhibits a higher minimum elevational limit of occupied habitat than either of the more apical sister species. This result is consistent with isolation of more persistent, peripheral populations at higher elevations, leading to speciation. Comparison of higher-elevation endemics to lower-elevation widespread species supports this interpretation. Such a finding affirms the importance of understanding geographic distribution on a scale appropriate to the action of vicariant mechanisms.

HISTORICAL BIOGEOGRAPHIC analysis has the use of Nelson and Platnick's (1981) concept become more rigorous because of the develop­ ofitems oferror to quantify the fit ofone pattern ment of the cladistic biogeographic method, in to another (Page 1990, 1993, 1994) have allowed which phylogenetic relationships of taxa are a more precise means to analyze congruence used as the basis for hypothesizing historical of biogeographic and coevolutionary patterns. relationships among areas of endemism defined Such methods have most often been aimed at by distributions of those taxa (Platnick and Nel­ illuminating patterns of vicariance, because the son 1978, Rosen 1978, Nelson and Platnick argument was made early on that dispersal pat­ 1981). Connecting the analysis of historical area terns were essentially random (Rosen 1978). relationships to historical relationships of taxa, More recently, the interpretation that both dis-' coupled with adoption of objective means to persal and vicariance can explain biogeographic hypothesize phylogenetic relationships (Hennig patterns, with such explanation dependent 1966, Farris et al. 1970) provided an objective on a hypothesis of phylogenetic relationships basis to the discipline. Recent developments in (Minaka 1987), eliminates the dichotomy be­ tween dispersal and vicariance explanations in biogeography, and replaces it with the search I Funding from the National Geographic Society (grant no. 4431-90) and the National Science Foundation (grant for common patterns and causes, versus distinct no. DEB-9208269) is gratefully acknowledged. This is con­ patterns and causes. tribution no. 1996-011 of the Hawai'i Biological Survey, The Hawaiian Islands are an ideal venue for Manuscript accepted 3 February 1997. exploration of such common and distinct pat­ 2 Department of Entomology, Comstock Hall, Cornell University, Ithaca, New York 14853-0901; Research Associ­ terns. Species-level diversity and endemicity are ate, B. P. Bishop Museum, Honolulu, Hawai'i 96817-0916, both extreme on the island chain, with as few 424 Dispersal and Vicariance in Hawaiian Carabids-LIEBHERR 425

as 230 to 250 ancestors necessary as founders goal worthy of all research programs, but espe­ for the over 5000 native (Zimmerman cially needed in Hawai'i because of the limited 1948, Nishida 1992, Miller and Eldredge 1995). geographic ranges of species and degradation Single-island endemicity approaches 100% in of native habitats due to invasion by exotics numerous taxa (Nishida 1992): for example, (Howarth 1985, Liebherr and Polhemus 1997). Ag1ycyderidae, Cerambycidae, and Curculioni­ dae (Coleoptera); Do1ichopodidae and Droso­ phi1idae (Diptera); Cixiidae and Delphacidae Hawaiian Platynini (Homoptera); Cosmopterygidae (Lepidoptera); The carabid tribe Platynini comprises Psocidae (Psocoptera). Because widespread dis­ over 2500 species in approximately 350 named tributions are anathema to cladistic biogeo­ genera, with an aggregate worldwide distribu­ graphic analysis (Nelson and Platnick 1981), tion. Species-level diversity and endemicity are these high levels of endemism permit derivation greatest in tropical montane forests, with major of area relationships from phylogenetic relation­ radiations in Mexico, Central America, and the ships in a relatively straightforward manner. Antilles (Whitehead 1973, Liebherr 1986, 1992), That much of the biogeographic pattern in South America (Moret 1990a,b, 1993, 1994), Hawai'i is based on overwater dispersal is indis­ Madagascar (Jeannel 1948, Basilewsky 1985), putable given current knowledge of the geologi­ Asia (e.g., Louwerens 1953), New Guinea (Dar­ cal history of the Islands (Clague and Dalrymple lington 1952), and the Hawaiian Islands (Sharp 1987, 1989). Nonetheless, the subsidence of 1903). The restricted geographic ranges ofmon­ Maui Nui affords the search for vicariant pat­ tane endemic species are due primarily to the terns among the resulting islands of Moloka'i, long-term duration of such habitats during cli­ Lana'i, and Maui. matic cycling, resulting in extensive adaptation This paper analyzes the biogeographic his­ of species to localized habitats (Southwood tory for a group of Hawaiian carabid beetles: 1977). This adaptation is best exemplified by the native species of the tribe Platynini. This the evolution of reduced flight capacity, through group composes one of only three tribes repre­ the shortening of the metathoracic episterna and sented in the native carabid beetle fauna, the flight wings in taxa of most montane tropical others being the tribe Bembidiini with 26 radiations ofthe tribe (e.g., Sharp 1903, Darling­ described species represented (Britton 1948a) ton 1952, Liebherr 1988). and the genus Mecyclothorax Sharp of the tribe The Hawaiian Platynini represent a uniquely Psydrini (Britton 1948b) with about 165 native prolific level ofdiversification, with 128 species species now known (J.K.L. and E. C. Zimmer­ found within the 16,640 km2 of the Islands man, unpubl. data). (Table 1). In comparison, Japan supports 140 The phylogenetic relationships of the native species (Habu 1978) within its area of 369,700 2 Platynini were used to derive a general hypothe­ km , Madagascar houses 326 species (Basilew- sis of area relationships (Liebherr and Zimmer­ man, in press). Working from that general TABLE 1 hypothesis, deviations including both dispersal­ based and vicariance-based elements were SPECIES DIVERSITY AND PERCENTAGE SINGLE-ISLAND ENDEMISM FOR HAWAIIAN PLATYNINI investigated. Discordant dispersal is objectively identified by items oferror criteria used to quan­ NO. NO. % tify the fit of observed c1adograms to particular ISLAND SPECIES ENDEMICS ENDEMISM patterns. Deviations from a general vicariance hypothesis for Maui Nui are explained by the Kaua'i 24 24 100 O'ahu 32 32 100 association of elevational shifts in peripheral Moloka'i 21 17 81 populations with speciation. Investigation of Llina'i 5 3 60 explanatory mechanisms beyond vicariance is West Maui 17 13 76 presented as a complementary approach to bio­ East Maui 35 31 89 Hawai'j 6 4 67 geographic analysis. Both methods require com­ Total 128 124 97 prehensive field collections with precise data: a 426 PACIFIC SCIENCE, Volume 51, October 1997

2 sky 1985) in 594,180 km , and New Guinea is the history ofa radiation resulting from coloniza­ home to 164 species (Darlington 1952, 1971) tion by a single common ancestor that has subse­ 2 over 808,510 km • Species distributions of quently been played out on all the high islands. Hawaiian platynines are extremely limited, with This not only adds another case study to the only four species found on more than one island growing core of studies dealing with biogeo­ (Table 1), and two of these widespread species graphic relationships of Hawaiian endemic restricted to the islands that formerly were part of organisms (Wagner and Funk 1995), but because the superisland Maui Nui. Even within islands, of the extreme endemicity and diversity of the known distributions may be limited to single group, also permits the search for repeated bio­ ridges or valleys. On Kaua'i, two undescribed geographic patterns among the included taxa. species ("atra" and "kahili") are known only Because cladistic biogeography attempts to sum­ from Mount Kahili, a southern spur of the marize all area relationships in a single clado­ Kawaikini-Wai'ale'ale central uplands. On gram of areas, deviations from that summary O'ahu, the 32 species include 8 restricted to the can be investigated in light of various attributes western Wai'anae range and 14 restricted to the of the attendant taxa. In this study, I first present eastern Ko'olau Mountains. the general biogeographic pattern exhibited by The extreme endemicity of the Hawaiian Hawaiian Platynini and then investigate concor­ platynines makes them ideal candidates for cla­ dant and discordant patterns of dispersal among distic biogeographic analysis (Platnick and Nel­ the major islands, as well as the vicariant pattern son 1978). Cladistic analysis of the Hawaiian among the fragments of the superisland Maui Platynini (Liebherr and Zimmerman, in press), Nui. based on 206 morphological characters and including all native Hawaiian species plus 41 Biogeographic Patterns outgroup taxa, indicates that the Hawaiian platynine radiation is monophyletic, with the The cladistic analysis of native Hawaiian closest outgroup the genus Lorostema Motschul­ Platynini (Liebherr and Zimmerman, in press) sky, distributed in India, Southeast Asia, New was used as the basis for this biogeographic Guinea and northern Australia, and the South­ analysis. The strict consensus cladogram of west Pacific as far as Tahiti (Figure 1). There­ 14,169 equally parsimonious cladograms-the fore, biogeographic analysis of the Hawaiian number of trees stored was limited by computer platynines provides the opportunity to deduce memory and the NONA tree-finding program rr Loros'ema (SE As;a, Japao, SW Padl~1 nsp. "mandibularis" (K)

I :'Pocaccus posticatus (K) UI Colpocaccus tantalus (0) [[Co/pocaceus ~naieosls (Mk + l + WM + EM)

Colpocaccus hawaiiensis (H)

FIGURE I. Cladistic relationships at base of Hawaiian platynine radiation and closest outgroup, Lorostema Motschulsky. Undescribed species name in quotation marks preceded by abbreviation "nsp." Area abbreviations are as follows: K, Kaua'i; 0, O'ahu; Mk, Moloka'i; L, Lana'i; WM, West Maui; EM, East Maui; H, Hawai'i (Big Island). Dispersal and Vicariance in Hawaiian Carabids-LIEBHERR 427

(Goloboff 1995)-showed that all discovered and the Big Island) and because the COMPO­ cladograms agreed on the monophyly of the NENT program of Page (1993) requires a fully group (Figure 1). Because the strict consensus resolved taxon-area cladogram, the first clado­ cladogram (e.g., Figures 2, 3) retained a consis­ gram stored in memory was arbitrarily chosen tent and repeated pattern of area relationships for this analysis. among the major islands (Kaua'i, O'ahu, Maui, When the resolved taxon cladogram was used Barypristus sharpi (EM) r----I Baryprislus incendiarius (H) Barypristus rupicola (EM) Colpodiscus lucipetens (Mk +L +WM +EM +H) nsp. kukui (H) Apteromesus maculatus (K) Mysticomenus mysticus (0) Mysticomenus tibialis (0) Chalcomenus costatus (K) Chalcomenus corruscus (0) Chalcomenus molokaiensis (Mk + WM +EM +H) nsp. polhemusi (Mk) nsp. pilikua (EM) r-----Atelothrus aaae (EM) Disenochus curtipes (K) nsp. polipoli (EM) Atrachycnemis sharpi (EM) Atrachycnemis perkinsi (Mk) Atrachycnemis koebelei (WM) Disenochus sulcipennis (K) nsp. atra (K) Disenochus erythropus (K) nsp. bryophila (K) Mauna frigida (EM) Disenochus munroi (Mk) nsp. viridis (WM) Disenochus aterrimus (K) Derobroscus micans (0) Brosconymus optatus (0) nsp. hihia (0) Anchonymus agonoides (EM) Disenochus longipes (Mk) nsp. pukalaina (WM) Disenochus fractus (EM) Disenochus agilis (EM) nsp. medeirosi (EM) Prodisenochus terebratus (EM) Broscomimus lentus (EM) Disenochus anomalus (EM) Disenochus brevipes (Mk) Disenochus cephalotes (WM) Disenochus micantipennis (K) nsp. waialeale (K) Anchotefflus elegans (K) Anchotefflus gracilis (0) Blackburnia insignis (0) Deropristus blaptoides (0) Deropristus punctatus (Mk) Deropristus deroderus (EM) nsp. kipahulu (EM)

FIGURE 2. Resolved cladogram of division I used for biogeographic analysis. Internal cladogram nodes to the right of stars collapse on the strict consensus cladogram. Undescribed species names preceded by abbreviation "nsp." Area abbreviations as in Figure 1. 428 PACIFIC SCIENCE, Volume 51, October 1997 K Metromenus aequalis (0) Metromenus fraternus (0) Melromenus limbatus (K) Metromenus pavidus (K) o Metromenus cuneipennis (0) Melromenus perpolilus (0) Metromenus meticulosus (0) Mk Atelolhrus fraclistriatus (0) nsp. fordi (0) nsp. foveolala (EM) Atelolhrus metromenoides (0) WM Metromenus hilaris (0) Metromenus caJigino511s (0) Metromenus mUlabilis (0) Metrornenus epicurus (0) EM nsp. hakeakapa (Mk) * nsp. kauwa (WM) nsp. fulgida (EM) l Atelothrus insociabilis (EM) Metromenus fraudator (Mk) Atelolhrus filipes (I.) H nsp. bartlelti (WM) Atelolhrus gracilis (WM) Alelothrus dyscoleus (EM) FIGURE 4. Single fundamental area cladogram that best Melromenus palmae (0) minimizes duplications, added leaves, and hypothesized Atelothrus platynoides (Mk) losses (Page 1993) for Hawaiian platynine carabid taxa. Area Metromenus sphodriformis (Mk +WM +EM) abbreviations as in Figure I. Atelolhrus depressus (I.) Metromenus moerens (Mk) Alelothrus longulus (WM) as the basis for cladistic biogeographic analysis ** Atelothrus constrictus (Mk) using items oferror as implemented in COMPO­ Atelothrus erro (EM) nsp. asquithi (K) NENT 2.0 (Page 1990, 1993), a single funda­ nsp. kahili (K) mental area cladogram (Figure 4) exhibited the nsp. huhula (0) Mesothriscus muscicola (0) best combination of fewest duplications, fewest nsp. paludicoJa (0) hypothesized losses, least items of error (Page Mesothriscus kauaiensis (K) Atelolhrus transiens (K) 1993), and strongest underlying character sup­ Mesothriscus optimus (K) port (Liebherr and Zimmerman, in press). This Mesothriscus alternans (K) hypothesis recognizes the dominant pattern of Mesothriscus opacus (K) Metromenus lenlus (H) area relationships exhibited by Hawaiian Mesothriscus lauaiensis (L) platynines to be progressive colonization from Mesothriscus vagans (Mk) nsp. auana (WM) Kaua'i to the younger islands. This progressive ambiens (K) pattern is at the level of superislands, as the Big Alelolhrus howarthi (EM) Island is the sister area to the islands previously Plalynus calalhifonnis (EM) Atelothrus debilis (Mk) united as Maui Nui: Moloka'i, Lana'i, and Maui. nsp. kuiki (EM) Therefore a vicariant pattern associated with the nsp. pauma (K) subsidence of Maui Nui (Clague and Dalrymple nsp. ulaula (EM) Mesothriscus Iricolor (Mk) 1987) is overlain on a progressive dispersal pat­ Mesolhriscus concolor (WM) tern composing distinct oceanic islands. The pre­ Mesosthriscus microps (WM) Metromenus latifrons (Mk) dominant pattern of Maui Nui vicariance Mesolhriscus abax (Mk) recognizes Moloka'i and West Maui as sister nsp. kolukala (EM) areas, with East Maui their sister area. Lana'i is nsp. abawides (EM) nsp. kuewa (EM) cladistically basal within Maui Nui. This general Mecumenus putealis (EM) pattern is at odds with the presence ofthe Pailolo MeCOSlomus perkinsi (EM) Mecomenus koebelei (WM) Metromenus audax (0) FIGURE 3. Resolved c1adogram of division 2 used for nsp. lihau (WM) Atelolhrus cheloniceps (Mk) biogeographic analysis. Internal c1adogram nodes to the right Metromenus calathoides (EM) of stars collapse on the strict consensus cladogram. Unde­ nsp. paloloensis (0) scribed species names preceded by abbreviation "nsp." Area Metromenus bardus (0) abbreviations as in Figure I. Melromenus oceanicus (0) Metromenus fossipennis (0) Metromenus fugitivtls (0) Metromenus prolervtls (0) Dispersal and Vicariance in Hawaiian Carabids-LIEBHERR 429

Channel between Moloka'i and Maui, and the Maui, suggesting that erosional aspects of the presence of an isthmus joining West and East Kohala Range may provide ecological require­ Maui. Under a strictly vicariant scenario, this ments necessary for this species' existence there. implies that mechanisms other than sea level The final two species-Colpodiscus lucipe­ inundation are responsible for fragmentation of tens (Blackburn) and undescribed species platynine habitats on Maui Nui; that is, vicariant "kukui"-are sisters and most parsimoniously patterns were established before final subsidence may be hypothesized to have been derived via of Maui Nui resulted in establishment of the allopatric speciation within the Big Island after PaHolo Channel. colonization of the Big Island by a common ancestor. Colpodiscus lucipetens is distributed across the island and on Maui Nui, whereas the Dispersal Patterns: Big Island Colonization undescribed species has been collected only in windward forest on the Big Island from Hilo to Based on the geologic history of the Islands KIlauea. The undescribed species is much less and the pattern ofarea relationships summarized common than C. lucipetens, and its known from all species, the most recent colonizations are likely to have occurred during the population restricted range may represent inadequate col­ ofthe Big Island. Based on the taxon-area clado­ lections from areas outside the historical collect­ gram, the six Big Island species have been ing axis along the Hilo-KIlauea Road (Manning derived from at least five colonization events 1986). Continued sampling of these taxa is (Table 2). Two of these involve sister species required to adequately assess their area of on Maui Nui: the sister pairs Colpocaccus sympatry. hawaiiensis Sharp and C. lanaiensis Sharp, and Based on the variety of divergence levels Barypristus incendiarius (Blackburn) and B. exemplified by colonization events among Big rupicola (Blackburn). In a third case, the Maui Island Platynini, the inescapable conclusion is Nui sister group comprises three species that the fauna of this island is being built by restricted to various islands, with Metromenus accretion of individually successful colonizing lentus Sharp now occurring on the Big Island. taxa. Major geologic events, such as massive Thus, colonization of the Big Island by M. lentus failures of older islands (Moore et al. 1989) may predates the origin of the three distinct species contribute to such colonizations, but synchro­ on Lana'i, Moloka'i, and West Maui. nous colonization does not appear to be the One Big Island species, Chalcomenus molo­ nonn. Three of the five colonizing episodes kaiensis Sharp, is widespread also on Maui Nui. involve species currently widespread across On the Big Island, C. molokaiensis is restricted islands that composed Maui Nui-Colpocaccus to the Kohala Mountains, the northernmost and hawaiiensis and C. lanaiensis, Chalcomenus oldest volcano composing the Big Island. The molokaiensis, and Colpodiscus lucipetens. Eco­ species is a streamside resident on Moloka'i and logical requirements for these species are poorly

TABLE 2

FLIGHT WING CONFIGURATION, SISTER TAXON, AND SISTER TAXON PROVENANCE OF SPECIES FOUND ON HAWAI'I (BIG ISLAND)

SPECIES WINGS SISTER TAXON PROVENANCE

CoLpocaccus hawaiiensis ++ CoLpocaccus Lanaiensis Maui Nui Barypristus incendiarius Barypristus rupicoLa East Maui CoLpodiscus Lucipetens ++ Undescribed sp. "kukui" Big Island Undescribed sp. "kukui" ++ CoLpodiscus Lucipetens Big Island ChaLcomenus moLokaiensis + ChaLcomenus moLokaiensis Maui Nui Metromenus Lemus Mesothrisclls Lanaiensis + M. vagans + undescribed sp. "auana" Maui Nui

NOTE: Macropterous metathoracic wings with wing ratio 3.0-5.0 (+ + J. macropterous wings with wing ratio 1.7-2.3 (+ J, vestigial wings (-J. 430 PACIFIC SCIENCE, Volume 51, October 1997

understood, but the association of the majority entire lineage from area A to area E leaving of colonizations with widespread species paral­ descendants in the intermediate areas. Such an lels the widespread distributions of colonists on approach is seriously deficient, because the pat­ Krakatau (Thornton et al. 1990). tern can also be explained by successive vicari­ Successful Big Island colonization has ance of a primordial distribution composed of occurred in both macropterous and brachypter­ all areas. Moreover, the simple example of five ous lineages (Table 2). Of the minimum five taxa in five areas with the relationship colonization events possible, three involve fully «A,B)(C(D,E))) precludes the ability to assign winged colonists and two involve vestigially a single unambiguous dispersal path connecting winged colonists. The flight-wing configuration the taxa in their areas. of the winged colonists varies. Both Colpocac­ Items of error analysis (Page 1990, 1994) cus hawaiiensis and Colpodiscus lucipetens pos­ permits use of an objective criterion to evaluate sess metathoracic wings 3.0--4.3 times the length how well a particular set of taxa adheres to a of the elytra (Liebherr and Zimmerman, in particular pattern of area relationships. In items press), the length hypothesized to be plesiomor­ of error analysis, an observed tree is modified phic for the Platynini. Records for these species through the addition of hypothetical terms so include captures on a window and in a window­ that its overall pattern of relationships is congru­ pane trap, respectively, suggesting that active ent with a particular general pattern. The hypo­ flight occurs in these species. Chalcomenus thetical terms are called added leaves (Page molokaiensis individuals possess flight wings 1993); the total number of leaves corresponds with full venation; however the alar surface is to the number of terminals in the modified only 1.7-2.3 times the elytrallength. All ecologi­ observed tree. Items of error are the difference cal information associated with this species in total cladogram edges between the modified involves collection in streamside habitats, under observed tree and the original observed tree. rocks, in litter, or on Cyanea plants. Though Because cladogram edges are two times the winged, flight is therefore not proven for this number of terminals in a dichotomous clado­ species. Both vestigially winged Big Island taxa gram, added leaves X 2 = items of error. (Barypristus incendiarius and Metromenus len­ As an example, the Colpocaccus clade (Fig­ tus) are sister to other vestigially winged taxa, ure 1) is perfectly congruent with the fundamen­ indicating that their differentiation on the Big tal area cladogram found for the entire platynine Island occurred in the absence of serviceable radiation (Figure 4). Colpocaccus cannot resolve flight wings. Successful colonization in the the Maui Nui areas, because C. lanaiensis is above cases therefore may occur by active flight, widespread across Moloka'i, Lana'i, West Maui, but as often by means not requiring flight, such and East Maui; however this lack of resolution as transport on floating trees or other occupied cannot conflict with the overall pattern. Other plant parts. parts of the resolved taxon-area cladogram fit less well. For example, the clade basally defined Older Dispersal Patterns by Disenochus aterrimus Sharp (Figure 2) de­ fines the area relationships (K(O(EM(Mk(WM, Hennig (1966) advanced the progression rule EM))))). To make this pattern congruent with to permit determination of historical dispersal the fundamental area cladogram (Figure 4), sev­ pathways. By this rule, taxa possessing the most eralleaves must be added: (1) Hawai'i must be derived character states are reasoned to occupy included as a sister area to Maui Nui, resulting the most recently colonized areas. Distributions in the addition of one leaf or two items of error; of common ancestors are determined by the dis­ (2) the redundant distribution of East Maui tribution of the patristically closest terminal requires the addition oftwo duplications and five taxa. He used the concept of rassenkreisen leaves so that the apical four taxa (in bold) are (sequences ofsubspecific taxa) to exemplify the present in three replications of an identical sub­ progression rule. Thus, a pectinate cladogram of pattern-«EM(Mk,WM), «EM(Mk,WM», infraspecific taxa inhabiting areas in the relation­ (EM(Mk,WM»)))-resulting in addition of five ship (A(B(C(D,E)))) indicates dispersal of the more terms or 10 items of error; and (3) the Dispersal and Vicariance in Hawaiian Carabids-LIEBHERR 431

missing area Uina'i must be inserted as sister gle terminals. This resulted in a total of 100 to the other Maui Nui areas, resulting in yet one terminals across the entire taxon-area cladogram more term (i.e., two more items of error). Thus, (Figures 1-3), a total within the capacity of the for the Disenochus aterrimus clade to fit the COMPONENT program (Page 1993). fundamental area cladogram (Figure 4) requires Exclusion of 37 of the 100 terminals resulted a total of seven additional leaves or 14 items of in no change in duplications, and exclusion of error. However, ifwe were to accept a hypothesis 61 others resulted in reduction by one duplica­ that dispersal by the ancestor ofD. fractus Sharp tion. A decrease of two duplications to a total into East Maui from West Maui occurred subse­ of58 was observed for only two terminals (Table quent to vicariant events congruent with the fun­ 3). Duplications can be viewed as speciation damental area cladogram, we could eliminate events undiscoverable because of the scale of the need for the five added leaves required under the areas ofendemism. Thus the speciation event point 2 above. This ad hoc assumption of dis­ might be considered "sympatric" at the level of persal can be viewed as equivalent to five added resolution of the areas of endemism, with both leaves, or 10 items of error. resultant taxa the common ancestors of their To assess the cost of assuming that relation­ own subsequently evolving lineage. Pectinately ships of each of the various taxa represent the redundant taxa (e.g., undescribed species "atra" general hypothesis of area relationships (Figure and Disenochus erythropus Sharp [Figure 2]) 4), each taxon was excluded sequentially from are one example of such a pattern. In the case calculation of duplications, leaves added, and of these two taxa, their redundancy requires losses based on the other taxa. The resultant insertion of leaves representing the other six totals were then compared with the totals based areas as sister to each species' representation. on all taxa (Table 3). The savings in fit criteria Thus a taxon exclusion resulting in only one were considered the decrease in duplications, fewer duplication is always associated with leaves, and losses relative to those of the entire redundancy in the cladogram. data set. These values were evaluated in the Exclusion of Metromenus palmae (Black­ context of savings accrued for all taxon exclu­ burn) results in reduction by two duplications sions; only the greatest relative savings merit (Table 3). Strong conflict with the fundamental acceptance of an explanatory ad hoc dispersal area cladogram explains this reduction (Figures hypothesis. In like manner, if a certain savings 3, 4). Metromenus palmae is the only O'ahu in the criteria merit acceptance of dispersal for taxon within an entire clade ofMaui Nui species. one particular taxon, then taxon exclusions The cladistically closest O'ahu species is Met­ resulting in greater savings must also merit romenus epicurus (Blackburn), eight nodes acceptance of dispersal. away. Assuming that M. palmae is the result of Before the analysis was conducted, mono­ back-dispersal from Maui Nui to O'ahu saves phyletic groups of taxa were condensed into sin- 23 added leaves, equivalent to 46 items of error. Those 23 added leaves are distributed in 18 TABLE 3 monophyletic sets, as indicated by the saving of 18 losses. No other taxon exclusion is even close FIT CRITERIA MAXIMIZED IN ITEMS OF ERROR ANALYSIS FOR ENTIRE TAXON-AREA CLADOGRAM AND FOR in such reductions of leaves added and losses CLADOGRAMS DERIVED AFTER EXCLUSION OF (Figures 5, 6). Therefore, if we assume that the FOUR SPECIES origin ofM. palmae involved dispersal ofa colo­ nizing propagule from the Moloka'i portion of LEAVES Maui Nui (Figure 3), we can reduce the leaves CLADOGRAM DUPLICATIONS ADDED LOSSES added by nearly 10% at a cost of one ad hoc All species 60 244 140 hypothesis of dispersal. Metromenus patmae 58 221 122 Excluding Disenochus fractus also results in Disellochus jractus 58 239 136 reduction by two duplications, and elimination Undescribed sp. of five added leaves and four losses (Table 3). "foveoJata" 60 229 135 Disenochus curtipes 60 237 133 This amount of reduction in leaves added is less than or equal to that observed because of 432 PACIFIC SCIENCE, Volume 51, October 1997

30

25

ctS 20 X ctS 15 r-. 0 10 Z 5 o• ill I I • I *ill 221 223 225 227 229 231 233235237239 241 243 245 Leaves Added

FIGURE 5. Distribution of numbers of leaves added for taxon exclusion trials. Analysis of all 100 terms of taxon-area cladogram requires 244 added leaves (*, see Table 3). exclusion of 38 other terminals (Figure 5); for associated with larger savings in items of error. losses the value is less than or equal to that We will return to this example when vicariant observed by exclusion of 14 other terminals patterns of Maui Nui are investigated later in (Figure 6). Therefore, savings in items of error the paper. are less than for numerous other taxa, and for It is important to analyze the cladistic rela­ us to invoke an ad hoc hypothesis of dispersal tionships and distributions oftaxa when evaluat­ in this case would call into question whether we ing the total reductions in items of error in such should consider it for the numerous other taxa an analysis. For example, exclusion of the unde-

30

25

20

15 * 5 o • • I 122 124 126 128 130 132 134 136 138 140 Losses

FIGURE 6. Distribution of losses for taxon exclusion trials. Analysis of all 100 terms of taxon-area cladogram requires 140 losses (*, see Table 3). Dispersal and Vicariance in Hawaiian Carabids-LIEBHERR 433

scribed species "foveolata" saves 15 added portion of extant suitable habitat for these leaves accounted for by five losses (Table 3), species. the first value falling well below the values of Viewing the five colonization events of the 98 other terminals (Figure 5). However, inspec­ Big Island in this more global (albeit islandic) tion of the taxon-area cladogram indicates that context, it appears that this youngest island has this savings reflects paraphyletic redundancy of received about 25% of the number of colonizing three taxa on O'ahu that are successive sister lineages observed for Maui Nui and those possi­ groups of the new species (Figure 3). In fact, ble for O'ahu. The isolation of Kaua'i is pro­ the pattern expressed is congruent with the fun­ found, because it has witnessed the emigration damental area cladogram in that a Maui Nui of propagules to O'ahu and/or Maui Nui, but taxon is related to those on O'ahu. has not yet delivered evidence of receiving a Disenochus curtipes Sharp and its four rela­ colonist from one of the younger islands. tives are hypothesized to be the sister group of the East Maui cave species Atelothrus aaae Samuelson & Liebherr (Figure 2). This relation­ Maui Nui Vicariance ship is supported by three characters: (1) reduced outer curvature of the eyes, (2) sparsely pubes­ One ofthe central tenets ofcladistic biogeog­ cent third antennomere, (3) elytral striae evenly raphy is the assumption of a single history of punctured throughout length (Liebherr and Zim­ areas decipherable from phylogenetic informa­ merman, in press). The first and second traits tion held in common among a variety of taxa are features often observed in cave species (Barr (Platnick and Nelson 1978). That there is a single 1983), and the third is reversed from the derived history is tautological. That it is decipherable condition wherein punctations are stronger on remains a challenge, with numerous techniques the base of the elytra than near the apex. Based advanced that provide different hypotheses of on the weak character support for its current area relationships given the same input of data placement, and biogeographic relationships of (Morrone and Carpenter 1994). The approach D. curtipes and its sister group, it seems unlikely taken here is slightly different. Rather than com­ that the current cladistic hypothesis is correct. pare methods of analysis, the fundamental area Be that as it may, the exclusion of D. curtipes cladogram (Figure 4) is used to identify patterns results in the second fewest losses (Figure 6), for taxa that conform and conflict with the gen­ but no reduction in duplications and only seven eral pattern of vicariance for Maui Nui. The fewer leaves added, a value equal to or less goal is to find distributional attributes other than than that for eight other terminals. Therefore, geographic range that might explain why a par­ acceptance of an ad hoc dispersal hypothesis ticular taxon deviates from the general pattern, whereby D. curtipes colonized Kaua'i from which is simply the predominant pattern in the Maui is not called for in this instance. taxon-area cladogram. Taking the fundamental area cladogram (Fig­ Six three-taxon triplets involving areas com­ ure 4) as the template, the taxon-area cladogram posing Maui Nui (Figure 8B-G) are implicit in was scanned and island transformations (i.e., the strict consensus cladogram (Liebherr and colonization events) noted (Figure 7). The single Zimmerman, in press). A seventh triplet is com­ back-dispersal event involving Metromenus pal­ posed of Disenochus munroi Perkins and unde­ mae is the lone exception to progressive coloni­ scribed species "viridis," sister taxa on the strict zation from older to younger islands. The 10 consensus (Figure 2), and Maunajrigida (Black­ colonization events linking Kaua'i and Maui Nui burn), sister group to these taxa in one class of may very well represent colonizations of O'ahu, most parsimonious cladograms, but only patris­ with subsequent extinction of the O'ahu species tically closest in the second arrangement ofthese on that highly eroded island. Currently, species. In the second arrangement, M. jrigida platynines are restricted to above 700 m on composes the outgroup to the resolved Anchony­ O'ahu, reducing the Ko'olaus to an extremely mus agonoides Sharp clade (Figure 2). The thin strip of suitable habitat. In the Wai 'anaes, monophyletic interpretation of M. jrigida, D. Mount Ka'ala represents the only substantial munroi, and the undescribed species is illustrated 434 PACIFIC SCIENCE, Volume 51, October 1997

FIGURE 7. Dispersal pathways derived from taxon-area cladogram using the fundamental area cladogram (Figure 4) as a template. Dispersals from Kaua'i to Maui Nui (area inside dashed lines) hypothesized to include colonization ofO'ahu based on general area relationships of fundamental area cladogram. Back-dispersal from Maui Nui to O'ahu supported by savings in items of error through taxon exclusion trials. for this analysis (Figure 8A), with interpretation West Maui undescribed species are found in under the alternate topology also discussed later. closed canopy to fragmented 'ohi'a forest at Three of these triplets (Figure 8A-C) are con­ lower elevations. From what we know of the gruent with the fundamental area cladogram habitat preferences for the other species, all are without addition of any leaves. Four others associated with mesic to wet 'ohi'a forest. The require added leaves to gain congruence, members of the Mesothriscus lanaiensis­ because ofsister-area relationships hypothesized vagans-undescribed species "auana" triplet (Fig­ for Moloka'i and East Maui (Figure 8D), Molo­ ure 8G) comprise streamside species; however ka'i and Lana'i (Figure 8£), West and East Maui they have been collected along streams in (Figure 8F), and Moloka'i and West Maui but 'ohi'a forest. lacking East Maui (Figure 8G). Therefore, The M. lanaiensis-vagans-undescribed spe­ among these seven triplets, five biogeographic cies "auana" triplet is also the single exception patterns are represented. to the elevational generalization that more basal A single unifying distributional attribute is groups are restricted to habitats at higher eleva­ predominant among the seven triplets. In six of tion. The basal species of this triplet, Mesothris­ the seven (Figure 8A-F), the basal species of the cus lanaiensis Sharp, is known from only two triplet has its lower elevationallimit ofhabitat at specimens collected by Perkins at the 2000-foot equal or higher elevation than either of the sister elevation on Lana'ihale in 1894 (Perkins 1894, species. In the case of Mauna frigida, this basal Sharp 1903), most likely in a streamside situa­ species occurs above timberline in open tion. More collections of species in this triplet, scrubland, whereas the Moloka'i D. munroi and and especially determination of whether the J~ frigida (EM; 1210) J~ sharpl (EM; 1210)

~munroi(Mk; 1040) ~ perkinsi (Mk; 1210)

A "viridis" (WM; 1210) B koebelei (WM; 910)

anomalus (EM; 1515) gracilis (WM; 700) ~L brevipes (Mk; 1180) platynoides (Mk; 575) cephalotes (WM; 890) dyscoleus (EM; 600)

Jc ;"",,~b;Ij, (EM; 910) longipes (Mk; 1150) --=~ fraudator (Mk; 820) "pukalaina" (WM; 1050)

E filipes (L; 605) fractus (EM; 1060)

J[; ~ I.",~",;, (L; 60s)

~[ vagans (Mk; 395)

G "auana" (WM; 1210)

sphodriformis (Mk; 910) molokaiensis (Mk; 820) =L sphodriformis (WM; 890) =L[ molokaiensis (WM; 600) H L sphodriformis (EM; 880) molokaiensis (EM; 700)

L/anaiensis (Mk; 100) [ lucipetens (Mk; 455)

~t lanaiensis (WM; 605) ~~ lucipetens (WM; 455)

~l,.".,en,;, (EM; 450) --:-ll:~"."' (E"; 150) J lanaiensis (L; 605) K lucipetens (L; 1020)

L constrictus (Mk; 910) cheloniceps (Mk; 1180) ~Ii== T longulus (WM; 910) "Iihau" (WM; 1270)

L erro (EM; 455) -;li.'==1= calathoides (EM; 1210)

FIGURE 8. A-G. Resolved triplets of single-island Maui Nui endemic species. Area abbreviations as in Figure I; lower elevational limit of known habitat in meters following area abbreviations; branch lengths for apical stems on c1adogram proportional to minimum elevational habitat limit. H-K. Lower elevational limits for widespread species on volcanoes composing the former Maui NuL L-M. Lower elevationallimits for ambiguous triplets of single-island Maui Nui endemic species. Undescribed species names in quotation marks. 436 PACIFIC SCIENCE, Volume 51, October 1997

West Maui undescribed species is found at eleva­ endemics whose relationships were ambiguous tions equivalent to those of M. vagans Sharp based on the strict consensus (Figure 8L, M). of Moloka'i, is the means to test the current Unlike the other resolved triplets (Figure 8A-G), elevational hypothesis. in these cases no single sister-species pair was Given the alternate topology for the Mauna represented in all most parsimonious clado­ jrigida triplet (Figure 8A), wherein Anchonymus grams. However, based on the elevational gener­ agonoides is the root of a clade sister to M. alization for the resolved triplets, it might be jrigida, the elevational generalization for this predicted that Atelothrus constrictus Sharp or triplet still holds. Anchonymus agonoides has A. longulus Sharp (Figure 8L) and undescribed been found no lower than 1210 m elevation, species "lihau" (Figure 8M) will be the cladisti­ a lower limit identical to that of M. jrigida. cally basal species in these two triplets after Therefore, no closely related species outside the more data are brought to bear. D. munroi-undescribed species "viridis" sister The failure to find a single vicariant pattern pair exhibits a lower elevationallimit than either for Maui Nui that fits all repetitions of areas sister species. across the taxon-area cladogram should not be This pattern is consistent with speciation via construed as a weakness of the cladistic biogeo­ peripheral isolation (Mayr 1963). Those periph­ graphie method. Without a cladistic hypothesis eral isolates that occupy higher elevations are of taxon relationships, no tests of any kind are the ones that persist longer as independent enti­ possible. Moreover, the presentation of a single ties and thereby are able to speciate. Isolated general pattern permits the search for deviants. populations at more climatically variable lower Extinction, quantified by items of error, can elevations: (1) have more opportunity to reestab­ account for disparities in how clades adhere to lish connections with other adjacent populations, the general pattern. The means to test this sort thus delaying speciation, or (2) have undergone of hypothesis include (1) showing that hypothe­ a higher rate ofextinction with or without specia­ sized "extinctions" really represent undiscov­ tion. By this scenario, speciation is the means ered taxa, and (2) generating more detailed by which morphological distinctiveness is fixed distributional data that might allow us to recast (Futuyma 1989). our definition of areas of endemism at a scale If higher-elevation populations are the ones appropriate to vicariant mechanisms. As shown isolated on various volcanoes, widespread spe­ above, extensive deviation from the general pat­ cies composing conspecific populations across tern costs numerous items of error, which can these same volcanoes should exhibit lower eleva­ be objectively mitigated through adoption of an tionallimits. In general this is so (Figure 8H-K). ad hoc dispersal hypothesis. Of the four widespread species, Colpocaccus Changing the scale ofareas ofendemism will lanaiensis and Colpodiscus lucipetens both occur cause us to define different numbers of wide­ as low as 100 to 150 m elevation at some sites. spread species and change the number of times Colpodiscus lucipetens is known from a single particular areas are redundantly represented on Lana'i specimen collected atop Lana'ihale (Fig­ the taxon-area cladogram. For example, the West ure 8K). Documenting permanent habitation of Maui mountains include the isolated block called Lana'i by this winged species requires subse­ LIhau that is topped with a forest no more than quent collections. Chalcomenus molokaiensis 0.65 km2 in area. This isolated forest is home occurs along streams and may have had a larger to seven species of Platynini, one of which is distribution with lower elevational limits before known only from this summit. Based on the the advent ofhuman introductions and deforesta­ definition of Platnick and Nelson (1978), this tion (e.g., Asquith and Messing 1993). One patch of forest could be defined as an area of might investigate infraspecific patterns of rela­ endemism. Recognizing LIhau in an analysis tionship in this case, predicting that Moloka'i would result in its placement seven times on the populations would be less closely related relative taxon-area cladogram, but the 10 other West to populations on West and East Maui (Figure Maui species would lack this area. Thus LIhau's 8/). relationships would be based on phylogenetic Two triplets are composed of single-island relationships of less than half the known West Dispersal and Vicariance in Hawaiian Carabids-LIEBHERR 437

Maui fauna. Moreover, the six LIhau species Island chain are more congruent than the vicari­ found at other sites would be treated as wide­ ant patterns on Maui Nui. This is in part because spread species: some distributed over discordant of the high cost established (i.e., relatively large sets of sampled localities on the volcano (e.g., numbers of items of error) for the adoption of Hana'ula, Pu'u Kukui, and 'Eke Crater). To date, an ad hoc dispersal hypothesis. Also, the areas a comparative approach whereby the scale of of endemism used to test dispersal are of greater endemism is modulated in repeated analyses of scale than the component volcanoes of Maui Nui the same aggregate area has not been done. used in the vicariance analysis. Given comprehensive sampling of Maui Nui The establishment of a biogeographic re­ localities, the rampant endemism of platynine search program that distinguishes common and carabids could allow such an attempt. The goal distinct patterns followed by a search for com­ would be to find an optimal mix of precise areas mon and distinct causation (Minaka 1987) is and informative endemism. Moreover, the stabil­ an advance over other methods that attempt to ity of hypotheses ofarea relationship throughout restrict causation before identifying patterns. such modulation should be investigated. Items of error analysis provides an objective Corroborating elevational habitat shifts as the means to accomplish this, permitting the integra­ overriding factor in the isolation and persistence tion of dispersal and vicariance, a goal pro­ of peripheral populations also requires a greater pounded by a variety ofbiogeographers (Croizat understanding of the distributional aspects of all 1962, Thornton 1983, Stace 1989, Grehan 1990, species. This scenario assumes that populational Briggs 1991). Nonetheless, a common dispersal processes operating at the level of individual pattern may still be based on independent dis­ demes work to produce shifts in occupied habi­ persal events, as shown for Big Island coloniza­ tat. Extinction of only a portion of species' tion by platynine carabids. Conversely, a populations may be the cause of discordant vic­ common vicariant history ofareas may be expe­ ariance. Therefore, a sound understanding of rienced differently by resident taxa facing popu­ how selection and limited effective population lational extinction and habitat alterations size can influence these species is essential. operating on a fine geographic scale. Bringing more character data to bear can improve our confidence in the phylogenetic rela­ tionships ofthe species, thereby eliminating mis­ takes in homology or polarity assessment as ACKNOWLEDGMENTS possible causes for disparate patterns. Corrobo­ ration using infraspecific divergence data elimi­ I thank Curtis Ewing, Zack Falin, Chris Mar­ nates the problem of extinct taxa, though it can shall, Michael McDonald, Jose Santisteban, never overcome the possibility that repeated Teresa Wells, and Kip Will, all of whom have bouts of gene exchange have replaced older pat­ worked on the Hawaiian carabid beetle project, terns of area relationship with newer ones. contributing their considerable talents to its Finally, addition of information from other taxa improvement. Adam Asquith, Curtis Ewing, Art endemic to the volcanoes of Maui Nui can test Medeiros, and Dan Polhemus have vastly both the general hypothesis of area relationships improved our knowledge of carabid diversity and the elevational generalization made here. and distribution. I thank Betsy Harrison-Gagne of the Hawai'i Natural Area Reserve System, Barrie Fox Morgan, Ed Misaki, and Mark White Conclusions of The Nature Conservancy of Hawai 'i, and Ed The diverse Hawaiian platynine species Petteys ofthe Hawai'i State Department ofLand swarm provides abundant information for gener­ and Natural Resources, for permission to work ating a general biogeographic hypothesis and in lands under their care. The Bishop Museum, then testing repeated congruent and discordant represented by Gordon Nishida, Scott Miller, patterns. Contrary to older arguments about the Dan Polhemus, and Al Samuelson, has sup­ random nature of dispersal (e.g., Rosen 1978), ported and assisted my activities in Hawai'i. I in this study the dispersal patterns across the thank Roderic Page and Rosemary Gillespie for 438 PACIFIC SCIENCE, Volume 51, October 1997 constructive reviews that helped improve the HARDT. 1970. A numerical approach to phylo­ manuscript. genetic systematics. Syst. Zool. 19:172-191. FUTUYMA, D. 1989. Speciational trends and the role of species in macroevolution. Am. Nat. 134:318-321. LITERATURE CITED GOLOBOFF, P. A 1995. NONA, version 1.5 [32 bit] [available from the author at Fundacion ASQUITH, A., and R. H. MESSING. 1993. Con­ Miguel Lillo, 205, 4000 San Miguel de Tucu­ temporary Hawaiian fauna of a low­ man, Argentina]. land agricultural area on Kaua'i: Implications GREHAN, J. 1990. Panbiogeography: Beyond for local and island-wide fruit fly eradication dispersal versus vicariance? 1. Biogeogr. programs. Pac. Sci. 47:1-16. 17:99-101. BARR, T. c., JR. 1983. The cavernicolous ancho­ HABu, A 1978. Fauna Japonica, Carabidae: menine beetles ofMexico (Coleoptera: Cara­ Platynini (Insecta: Coleoptera). Keigaku bidae: Agonini). Assoc. Mex. Cave Stud. Pub!. Co., Tokyo. Bull. 8: 161-192rrex. Mem. Mus. Bull. HENNIG, W. 1966. Phylogenetic systematics. 28:161-192. University of Illinois Press, Urbana. BASILEWSKY, p. 1985. Insectes Coleopteres, HOWARTH, F. G. 1985. Impacts of alien land Carabidae . Faune de Madagascar and mollusks on native plants and 64:543 p. in Hawaii. Pages 149-179 in C. P. BRIGGS, J. C. 1991. Historical biogeography: Stone and 1. M. Scott, eds. Hawaii's terrestrial The pedagogical problem. J. Biogeogr. ecosystems: Preservation and management. 18:3-6. University of Hawai'i Press, Honolulu. BRITTON, E. B. 1948a. The carabid tribes Har­ JEANNEL, R. 1948. Coleopteres carabiques de la palini, Lebiini, and Bembidiini in Hawaii. region Malgache (2e partie). Faune l'Emp. (Coleoptera). Proc. Hawaii. Entomol. Soc. Fr. 10:373-765. 13:235-254. LIEBHERR, J. K. 1986 (1987). A taxonomic revi­ ---. 1948b. A revision ofthe Hawaiian spe­ sion of the West Indian Platynus beetles cies of Mecyclothorax (Coleoptera: Carabi­ (Coleoptera: Carabidae). Trans. Am. Ento­ dae). Occas. Pap. Bernice Pauahi Bishop mol. Soc. (Phila.) 112:189-268. Mus. 19:107-166. ---. 1988. Biogeographic patterns of West CLAGUE, D. A, and G. B. DALRYMPLE. 1987. Indian Platynus beetles. Pages 121-152 in 1. The Hawaiian-Emperor volcanic chain, part K. Liebherr, ed. Zoogeography of Caribbean 1, geologic evolution. Pages 5-54 in R. W. insects. Cornell University Press, Ithaca, Decker, T. L. Wright, and P. H. Stauffer, eds. New York. Volcanism in Hawaii, Vol. 1. U.S. Geol. Surv. ---. 1992. Phylogeny and revision of the Prof. Pap. 1350. Platynus degallieri species group (Coleop­ ---. 1989. The Hawaiian-Emperor chain. tera: Carabidae: Platynini). Bull. Am. Mus. The geology of North America (The Eastern Nat. Hist. 214: 115 p. Pacific Ocean and Hawaii) N:187-287. LIEBHERR, J. K., and D. A POLHEMUS. 1997. CROIZAT, L. 1962. Space, time, form: The bio­ Comparisons to the century before: The leg­ logical synthesis. Published by the author, acy of R. C. L. Perkins and Fauna Hawai­ Caracas. iensis as the basis for a long-term ecological DARLINGTON, P. J., JR. 1952. The carabid beetles monitoring program. Pac. Sci. 51:490-504. of New Guinea. Part 2. The Agonini. Bull. LIEBHERR, J. K., and E. C. ZIMMERMAN. Cladis­ Mus. Compo Zool. 107:89-252 + 4 pIs. tic analysis, phylogeny, and biogeography of ---. 1971. The carabid beetles of New the Hawaiian Platynini (Coleoptera: Carabi­ Guinea. Part IV. General considerations; anal­ dae). Syst. Entomol. (in press). ysis and history of fauna; taxonomic supple­ LOUWERENS, C. 1. 1953. The Oriental species ment. Bull. Mus. Compo Zool. 142:129-337. of Mad. (Col. Carabidae). Treu­ FARRIS, 1. S., A. G. KLUGE, and M. 1. ECK- bia 22:75-151. Dispersal and Vicariance in Hawaiian Carabids-LIEBHERR 439

MANNING, A. 1986. The Sandwich Islands Com­ PAGE, R. D. M. 1990. Component analysis: A mittee, Bishop Museum, and R. C. L. Perkins: valiant failure? Cladistics 6: 119-136. Cooperative zoological exploration and pub­ --.1993. COMPONENT, version 2.0. The lication. Bishop Mus. Occas. Pap. 26: 1--46. Natural History Museum, London. MAYR, E. 1963. species and evolution. ---. 1994. Parallel phylogenies: Recon­ Harvard University Press, Cambridge, structing the history of host-parasite assem­ Massachusetts. blages. Cladistics 10:155-173. MILLER, S. E., and L. G. ELDREDGE. 1995. Num­ PERKINS, R. C. L. 1894. Diary of collecting on bers of Hawaiian species: Supplement 1. Lanai, 1894. Copies of ms. made by Perkins Bishop Mus. Occas. Pap. 45:8-17. in 1936, Bishop Museum archives, SC Per­ MINAKA, N. 1987. Branching diagrams in cla­ kins, Box 1.3. Copies also in Entomology distics: Their definitions and implications for Library, The Natural History Museum, biogeographic analysis. Bull. Biogeogr. Soc. London. Jpn.42:1-1O. PLATNICK, N., and G. NELSON. 1978. A method MOORE, 1. G., D. A. CLAGUE, R. T. HOLCOMB, for analysis of historical biogeography. Syst. P. W. LIPMAN, W. R. NORMARK, and M. E. Zool. 27:1-16. TORRESAN. 1989. Prodigious submarine ROSEN, D. E. 1978. Vicariant patterns and histor­ landslides on the Hawaiian Ridge. J. Geo­ ical explanation in biogeography. Syst. phys. Res. 94(B12): 17,465-17,484. Zool. 27:159-188. MORET, P. 1990a. Demembrement du genre Col­ SHARP, D. 1903. Coleoptera, 2. Pages 175-292 podes auctorum. II. genre Neotropical Dys­ in Fr. ..ma Hawaiiensis, Vol. 3, part 3. Cam­ colidion n. gen. (Coleoptera, Caraboidea, bridge University Press, Cambridge. Platyninae). Rev. Fr. Entomol. (n.s.) SOUTHWOOD, T. R. E. 1977. Habitat, the templet 12:67-80. for ecological strategies? J. Anim. Ecol. ---. 1990b. Les Dyscolus equatoriens du 46:337-365. groupe bucculentus sp. nov. (Coleoptera, Ca­ STACE, C. A. 1989. Dispersal versus vicari­ rabidae). Nouv. Rev. Entomol. (n.s.) ance-no contest! 1. Biogeogr. 16:201-202. 7:271-281. THORNTON, I. W. B. 1983. Vicariance and dis­ ---. 1993. Les Dyscolus de1,Equateur: Revi­ persal: Confrontation or compatibility? Geo­ sion des especes aelytres achetes (lere partie) Journal 7.6:557-564. (Coleoptera, Harpalidae, Platyninae). Rev. Fr. THORNTON, I. W. B., T. R. NEW, R. A. ZANN, Entomol. (n.s.) 15:1-13. and P. A. RAWLINSON. 1990. Colonization of ---. 1994. Les Dyscolus de l'Equateur: Revi­ the Krakatau Islands by animals: A perspec­ sion des especes aelytres achetes (2e partie) tive from the 1980s. Philos. Trans. R. Soc. (Coleoptera, Harpalidae, Platyninae). Rev. Fr. Lond. B BioI. Sci. 328:131-165. Entomol. (n.s.) 16:1-11. WAGNER, W. L., and V. A. FUNK, eds. 1995. MORRONE, 1., and 1. M. CARPENTER. 1994. In Hawaiian biogeography: Evolution on a hot search of a method for cladistic biogeogra­ spot archipelago. Smithsonian Institution phy: An empirical comparison of component Press, Washington. analysis, Brooks parsimony analysis, and WHITEHEAD, D. R. 1973. Annotated key to three-area statements. Cladistics 10:99-153. Platynus, including Mexisphodrus and most NELSON, G., and N. PLATNICK. 1981. Systemat­ "Colpodes," so far described from North ics and biogeography, cladistics and vicari­ America including Mexico (Coleoptera: Ca­ ance. Columbia University Press, New York. rabidae: Agonini). Quaest. Entomol. 9: NISHIDA, G. M. 1992. Hawaiian terrestrial 173-217. checklist. Hawai'i Biological Sur­ ZIMMERMAN, E. C. 1948. Insects ofHawaii, Vol. vey, Honolulu. 1. University of Hawai'i Press, Honolulu.