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Blackburnia kavanaughi, new species, previously misidentified as Blackburnia asquithi Liebherr, is described from the western portion of the Alakai Plateau, Kauai, Hawaii. The species is placed phylogenetically based on cladistic analysis including all known Hawaiian Blackburnia, supporting membership in the subgenus Metromenus Sharp. The new species is hypothesized to be most closely related to B. debilis (Perkins) of Molokai and B. kuiki Liebherr of East Maui, sharing enhanced pubescence of the antennal pedicel, reduction in number of the dorsal elytral setae, and elongate ventrolateral setae of tarsomere 5. Individuals of all three species reside in deep arboreal moss mats associated with ohia lehua trees (Metrosideros polymorpha: Myrtaceac). Blackburnia debilis, previously known only from the single male holotype collected in 1902, was recollected in 2005 from Uapa summit, Molokai, in arboreal moss on the mesic leeward edge of ohia lehua forest. Discovery of B. kavanaughi on Kauai implies an additional colonization event from Kauai to newer islands during the diversification of Blackburnia, while corroborating the previous general biogeographic pattern of progressive colonization of the Hawaiian Islands by this clade. Phylogenetic placement of B. debilis + B. kuiki as adelphotaxon to B. kavanaughi results in a hypothesized sister-group relationship between sympatric sister species-the epigean B. calathiformis (Sharp) and the troglobitic B. howarthi (Samuelson and Liebherr)-corroborating F. G. Howarth's general hypothesis for the evolution of Hawaiian troglobites.
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Author(s): James K. Liebherr
Source: Journal of the New York Entomological Society,
Published By: The New York Entomological Society
DOI: http://[17:RADOBK]2.0.CO;2
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Department of Entomology, Comstock Hall, Cornell University,
Ithaca, New York 14853-2601, U.S.A.
Abstract.—Blackburnia kavanaughi, new species, previously misidentified as Blackburnia asquithi
Liebherr, is described from the western portion of the Alakai Plateau, Kauai, Hawaii. The species is
placed phylogenetically based on cladistic analysis including all known Hawaiian Blackburnia,
supporting membership in the subgenus Metromenus Sharp. The new species is hypothesized to be
most closely related to B. debilis (Perkins) of Molokai and B. kuiki Liebherr of East Maui, sharing
enhanced pubescence of the antennal pedicel, reduction in number of the dorsal elytral setae, and
elongate ventrolateral setae of tarsomere 5. Individuals of all three species reside in deep arboreal
moss mats associated with ohia lehua trees (Metrosideros polymorpha: Myrtaceae). Blackburnia
debilis, previously known only from the single male holotype collected in 1902, was recollected in
2005 from Uapa summit, Molokai, in arboreal moss on the mesic leeward edge of ohia lehua forest.
Discovery of B. kavanaughi on Kauai implies an additional colonization event from Kauai to newer
islands during the diversification of Blackburnia, while corroborating the previous general
biogeographic pattern of progressive colonization of the Hawaiian Islands by this clade.
Phylogenetic placement of B. debilis +B. kuiki as adelphotaxon to B. kavanaughi results in
a hypothesized sister-group relationship between sympatric sister species—the epigean B.
calathiformis (Sharp) and the troglobitic B. howarthi (Samuelson and Liebherr)—corroborating
F. G. Howarth’s general hypothesis for the evolution of Hawaiian troglobites.
Key words: adaptation, phylogeny, biogeography, speciation.
As explicit representations of character congruence and incongruence supporting
a hierarchical network of terminals, cladistic hypotheses can be reevaluated after
discovery of any new character or taxon that necessarily expands knowledge concerning
the focal clade. The Hawaiian Blackburnia beetles have afforded an empirical example
of such activities, as continued field survey has repeatedly uncovered previously
uncharacterized taxa (Liebherr and Zimmerman 2000, Liebherr 2001, 2003, Liebherr
and Short 2006). In this paper, I present information characterizing a species previously
collected in 1991 on the island of Kauai, but at that time confused with another more
common species, Blackburnia asquithi Liebherr (Liebherr and Zimmerman 2000). This
previous confusion was brought to light through a 2005 field trip to Kauai, where
additional specimens of this new species were collected from moss mats on the mesic
edge of the Alakai Swamp. A subsequent search of Blackburnia specimens collected by
David H. Kavanaugh, California Academy of Sciences uncovered several more
specimens, with the aggregate set of taxonomic material sufficient to support a revised
phylogenetic analysis
In this paper I first describe the new species, and then present a revised phylogenetic
hypothesis based on morphological characters. Inclusion of the new species slightly
J. New York Entomol. Soc. 114(1–2):17–27, 2006
modifies the cladistic structure of subgenus Metromenus Sharp of Hawaiian Black-
burnia, to which the new species belongs. Ramifications of this change to the
phylogenetic hypothesis include corroboration of the general biogeographic pattern for
this group, and reassessment of the evolutionary origin of a cave-inhabiting troglobite.
Specimen Collection, Description and Deposition.
Specimens were collected by applying synthetic pyrethrin insecticide to arboreal moss
mats, with beetles and other insects emerging from the moss to fall onto a white sheet
placed below. Nighttime searching with headlamps allowed collection of adults walking
on the ground.
Standardized body length equals the sum of: 1, median head length from labral margin
to cervical ridge; 2, median pronotal length; and 3, the distance from scutellar base to the
apex of the left elytron. A 0.1 mm resolution millimeter rule was used to calibrate the
ocular grid used to measure lengths. The ocular ratio is defined as maximum width across
the compound eyes, measured from above, divided by the minimum width of the frons
measured between the eyes (Liebherr and Zimmerman 2000).
Terminology used to describe male genitalia and female reproductive tract structures
follow Liebherr and Will (1998) and Liebherr and Zimmerman (2000). Aedeagal
structures are described using the anatomically plesiomorphic orientation, i.e. the point
of parameral articulation is on the euventral surface of the median lobe (Sharp and
Muir 1912), with other surfaces described relative to that position.
Type specimens are deposited in the Bernice Pauahi Bishop Musuem, Honolulu
(BPBM), California Academy of Sciences, San Francisco (CAS), and Cornell University
Insect Collection, Ithaca (CUIC).
Cladistic Analysis. The character-taxon matrix first published by Liebherr and
Zimmerman (1998) and subsequently modified through addition of this species
plus three others (Liebherr 2001, 2003, Liebherr and Short 2006) was analyzed cladis-
tically. The complete matrix is available at
Analytical protocols are identical to those used in Liebherr (2003), with the exception
that the search was limited to 12 ratchet runs (Nixon 1999, 2002) of 1000 iterations each,
with those trees entered to Nona (Goloboff 1999) for max* branch swapping. Bremer
(1994) support was calculated for 1000 suboptimal trees each of 1–10 steps longer than
the most parsimonious trees. One of the multiple equally parsimonious trees (MEPT)
was chosen for presentation, but Bremer supports of 0 in the figure indicate nodes that
were collapsed in the strict consensus of all MEPTs. This paper presents the current
preferred cladogram of subgenus Metromenus Sharp, which is the sister group to
subgenus Blackburnia Sharp. The current preferred cladogram for subgenus Blackburnia
is presented in Liebherr and Short (2006).
Blackburnia (Metromenus)kavanaughi, new species
Blackburnia (Metromenus)asquithi Liebherr, Liebherr and Zimmerman 2000: 263 (in
part, misidentification).
Diagnosis. Eyes reduced, ocular ratio 1.25–1.33 (Fig. 1); pedicel with 3–5 elongate setae
on outer surface; pronotal lateral margins glabrous, basolateral margins concave
anterad obtusely rounded hind angles, basal margin trisinuate, the margins convex
posterad laterobasal depressions; elytral humeri obtusely angulate; dorsal elytral setae 2
or 1 each side, either anterior seta or anterior plus middle setae absent; basal
metatarsomeres 1–3 medially sulcate, metatarsomere 5 with 4–6 evident ventrolateral
setae subequal in length to tarsomere depth at point of setal insertion; metathoracic
flight wings vestigial, lobelike flaps very short, not extended beyond hind margin of
metathorax; head capsule dark brunneous, pronotum brunneous with lateral margins
rufoflavous, elytral disc shiny piceous with sutural intervals, base and lateral margins
anterad subapical sinuation flavous; standardized body length 6.1–6.7 mm. Female
specimens have three setae each side of apical abdominal ventrite. Within subgenus
Metromenus—where this species is placed based on the dorsally sulcate basal tarsomeres
and vestigial hind wings—reduced eyes are also observed in B. debilis (Perkins) of
Molokai and B. howarthi (Samuelson and Liebherr) of East Maui. The new species
shares elongate setae of the antennal pedicel and absence of the anterior dorsal elytral
setae with B. debilis and B. kuiki Liebherr of East Maui. The glabrous pronotal margins
are also shared with those two species, as well as with numerous species in Oahu
(Liebherr and Zimmerman 2000).
Description. Head. Frons and vertex broadly convex due to extremely reduced eyes,
anterior supraorbital seta in well-developed fovea, but frontal impressions almost
absent, traceable only as irregular wrinkles extended from frontoclypeal suture;
compound eye with approximately 15 ommatidia aligned across horizontal diameter;
neck with broad, distinct dorsal transverse impression; antennae elongate, extended
almost to midpoint of elytra; labrum with anterior margin shallowly and broadly
emarginate, six setae present but medial pair shorter than lateral neighbors; mentum
tooth broad, slightly bifid apically.
Prothorax. Pronotum broadest before middle of length measured along midline,
lateral marginal depressions broad, shallow, lateral margin upraised but without
marginal bead, margin broader posterad basal sinuation, this broader margin extended
to form raised hind margin of laterobasal depressions; laterobasal depressions smooth,
with small median tubercle, deepest linearly along mesal margin; median base
impunctate, unmargined; median longitudinal impression shallow, slightly deeper near
shallow and smooth anterior transverse impressions; anterior marginal bead continuous
though fine, deepened near rounded, moderately projecting front angles. Prosternal
process unmargined, flat ventrally, glabrous; prosternum with shallow vertical
impression about 1/5 of length from anterior margin.
Elytra convex apically, widest just behind middle measured along midline, narrowed
anteriorly with outer margin rounded at humeri; parascutellar setae present; dorsal elytral
setae (middle or posterior setae when present) set in inconspicuous depressions associated
with second stria (though one specimen with two setae on one elytron has posterior seta
associated with constriction of second interval; CAS); elytral striae smooth, continuous,
evident to apex, with slight irregularities in basal half of elytral length; elytral intervals
moderately convex; lateral margins broad from position of first lateral elytral seta to
subapical sinuation, which is distinguished as an abrupt change in curvature of the elytral
margin, resulting in a subtruncate elytral apex; basal groove distinctly angulate at
humerus, where basal groove meets lateral elytral margin; 12–14 lateral elytral setae.
Pterothorax. Metepisternum appearing slightly elongate, length of mesal margin 1.13
length of anterior margin.
Legs. Profemur with two posteroventral setae; mesofemur wth two anteroventral
setae; metacoxa bisetose, inner seta absent; metafemur dorsoapically glabrous;
tarsomeres broadened apically; basal metatarsomere stout, less than 23length
of mesal metotibial spur; apical setae of metatarsomere 4 positioned subapically
on outer edge of tarsal lobe, the lobes very short, tarsomere apex only slightly
Coloration. Palpi, antennae, coxae, trochanters and femora testaceous, contrastedly
paler than body; femora rufotestaceous, dorsolateral and dorsal carinae darker,
brunneous, tarsomeres similarly brunneous matching pronotal and elytral epipleura;
thoracic and abdominal ventrites slightly darker than femora and tarsomeres, matching
piceous elytral disc.
Microsculpture. Vertex shiny but with evident isodiametric sculpticells, the mesh
stretched transversely in dorsal impression of neck; pronotal lateral depressions,
laterobasal depressions, and median base with well-developed isodiametric micro-
scuplture, disc with well-developed transverse mesh microsculpture; elytral intervals
with well-developed miscrosculpture of transversely stretched isodiametric sculpticells in
rows, sculpticell margins shallower at elytral apex, the surface shinier; thoracic and
abdominal ventrites with transverse mesh microsculpture, the surface appearing
granulate due to the distinct sculpticell margins.
Male genitalia. Aedeagal median lobe pale, gracile, depth approximately 0.203
distance from parameral articulations to tip (Fig. 2); euventral surface straight medially,
ventral excavation distinct; median lobe apex downturned, length of tip extended
beyond insertion of the internal sac approximately equal to tip depth at sac ostium;
sagittal crest present; median lobe apex in ventral view tightly rounded, the left margin
straight for a short distance before tip (Fig. 3); internal sac lightly sclerotized,
microspicules evident only due to folds of the inverted sac.
Female reproductive tract. Gonocoxa broad; gonocoxite 1 with apical fringe of setae,
6–8 setae in three rows at medioapical angle, 2–3 setae laterally (Fig. 4); gonocoxite 2
broad basally with narrowly rounded apex, bearing two lateral ensiform setae and one
dorsal ensiform seta; apical nematiform setae short, 0.163length of gonocoxite 2; bursa
copulatrix symmetrical, distance from base to spermathecal duct subequal to length of
common oviduct (Fig. 5); bursa with medial band of spikelike bursal microtrichia, and
dorsal bursal pouch lined with granulate microsculpture; spermatheca ovoid, reservoir
about 1/3 length of spermathecal duct with fewer than 20 taenidia-like constrictions
along length; spermathecal gland entering at reservoir-spermathecal duct junction.
Types. Holotype, male, (Fig. 1), HI: Kauai NaPali-Kona For. Res. Alakai Swamp Tr.
14-V-1991 el. 1150-1180 m J.K. Liebherr on trail at night//Paratype Blackburnia
(Metromenus) asquithi Liebherr 2000 (yellow paper)//PARATYPE Cornell U.
No. 7122//HOLOTYPE Blackburnia (Metromenus) kavanaughi Liebherr 2006 (red
paper) (CUIC). Paratypes: same data as holotype (CUIC, 1 male); HI: Kauai, NaPali-
Kona For. Res., Jct. Alakai Swamp & Pihea Trails, 1190 m, 14-V-1991, stop #91-26,
D.H. Kavanaugh (CAS, 2 males); same data but 16-V-1991, stop #91-29 (CAS, 1
female); HI: Kauai Napali-Kona For. Res. Alakai Swamp Pihea Tr. Jct. E, 1205m,
22u07.809N 159u37.669W, 17-V-2005 lot 02, pyrethrin fog mossy ohia +Ilex, J.K.
Liebherr (BPBM, CUIC, 2 females); HI: Kauai, Alakai Swamp Tr. E Kawaikoi Str., el.
1230 m, 22u08.859N 159u36.399W, 20-V-2005 lot 03, pyrethrin fog mossy ohia log, J.K.
Liebherr (CUIC, 1 female).
Etymology. The patronymic species epithet kavanaughi honors fellow carabidologist
David H. Kavanaugh, who collected this species when it was first encountered in 1991.
Distribution. This species is known only from the northwest portion of the Alakai
Plateau, in forests bordering Kawaikoi Stream (Fig. 6).
Habits. Individuals of this species have been found running on the ground at night on
muddy portions of a trail, and by applying pyrethrin insecticide to mossy surfaces of
ohia lehua (Metrosideros polymorpha: Myrtaceae) and Hawaiian holly (Ilex anomala:
Aquifoleaceae) growing on nurse logs of ohia lehua. The reduced eyes of the adult are
consistent with habitation in deep moss (Liebherr and Short 2006).
Figs. 1–3. Blackburnia kavanaughi, n. sp. 1. Male holotype, dorsal view. 2. Male aedeagal
median lobe, right lateral view (exerted position); ve, medioventral excavation of aedeagal median
lobe. 3. Male aedagal median lobe apex, euventral view.
Identification. ‘‘Key 4, Adults of subgenus Metromenus Sharp, Division 2 (Sharp 1903),
excluding subgenus Colpocaccus Sharp (Liebherr and Zimmerman 2000, p. 66)’’ can be
modified by insertion of one and renumbering a second couplet to permit identification
of B. kavanaughi, new species:
65(64). Eyes convex, outer surface more convex than curvedefined by gena, ocular ratio 1.45–1.58;
elytra with 3 dorsal elytral setae each side, apex broadly paler than disc . . . 65a
Eyes much less convex, outer surface curved with same radius as gena, ocular ratio
1.2–1.33; elytra with 1–2 dorsal elytral setae, the anterior always absent, the second
setal position polymorphic, elytral apex concolorous with disc, though sides and
base paler; Kauai ....................... Blackburnia kavanaughi, new species
65a(65). Elytral intervals 4–6 markedly convex basally, subcarinate; elytra with distinctly de-
veloped isodiametric sculpticells arranged in transverse rows; Kauai . . ........
................................... Blackburnia limbata (Sharp) (p. 404)
Elytral intervals 4–6 flat basally, striae obsolete on elytral base; elytra with micro-
sculpture of distinctly transverse sculpticells; Oahu, Waianae Range . . ........
................................. Blackburnia hilaris (Blackburn) (p. 377)
Blackburnia debilis (Perkins)
Discussion. Blackburnia debilis was described by Perkins (1917) from a single male
specimen labeled ‘‘Molokai 3000 ft. iii1902 RCLP.’’ Based on Perkins’ notes and
activities, this label was interpreted to indicate the vicinity of his 3000 ft. elevation
Makakupaia base camp (Liebherr and Zimmerman 2000). As this area is currently
extensively afforested with alien conifers, the continued existence of B. debilis was
therefore questioned (Liebherr 2004), and this species was proposed as one of two
Molokai Blackburnia to have suffered anthropogenic extinction.
On 11–12 May 2005, an insect survey was conducted along the newly renovated bird
transect running from Puu Haha (1100 m elevation) to Uapa summit (1500 m
elevation), a small prominence on the Kamakou Volcano rim just west of the highest
point, Mt. Kamakou. A single male specimen of B. debilis was collected just above the
ecotonal transition from uluhe (Dicranopteris linearis: Gleicheniaceae) fern-dominated
habitat to mesic ohia woodland. Label data are: ‘‘HI: Molokai, Puu Haha-Uapa
transect, 12-V-2005 lot 01, 21u06.249N 156u52.419W, el. 1355 m, pyrethrin fog mossy
ohia, J.K. Liebherr (CUIC).’’ The collecting site was within one of the lowest elevation
Figs. 4–6. Blackburnia kavanaughi, n. sp. 4. Female left gonocoxa, ventral view. 5. Female
reproductive tract, ventral view. Codens include: afs, apical fringe of setae, basal gonocoxite; ans,
apical nematiform setae; bc, bursa copulatrix; co, common oviduct; des, dorsal ensiform setae, dp,
dorsal pouch of bursa copulatrix; gc1, basal gonocoxite; gc2, apical gonocoxite; les, lateral ensiform
setae; sg, spermathecal gland; sp, spermatheca. 6. Distributional records. Figure 7. Strict consensus
of seven equally parsimonious cladograms for species of subgenus Metromenus; 1118-step length,
CI 50.16, RI 50.72 (see text). Arrow indicates 1-step cost associated with moving B. debilis +B
kuiki to previously hypothesized position as adelphotaxon to B. howarthi (Liebherr 2003). Island
codens include: K, Kauai; O, Oahu; Mk, Molokai; L, Lanai; WM, West Maui; EM, East Maui; H,
Hawaii Island.
copses of ohia lehua trees with moist mossmats, and was considerably drier than the
habitats at the summit. The pyrethrin fog treatment was applied at approximately
0830 h while hiking up the ridge, and then checked on the way down at 1600 h.
Accompanying the lone B. debilis in the sample were B. abax (Sharp) (1, CUIC), B.
constricta (2, CUIC) and B. platynoides (1, CUIC). Of these, B. platynoides is restricted
to lower-elevation habitats, where individuals have been most commonly found in the
leaf axils of the monocotyledonous ieie (Freycinetia arborea: Pandanaceae) (Liebherr
and Zimmerman 2000).
Cladistic analysis recovered 10,108 cladograms of 1112-step length (CI 50.16, RI 5
0.72), and in all equally parsimonious trees B. kavanaughi was unambiguously placed as
the adelphotaxon to the species pair B. debilis of Molokai plus B. kuiki of Haleakala,
East Maui within subgenus Metromenus of genus Blackburnia (Fig. 7). Other relation-
ships across Blackburnia were more ambiguous, with alternate placements of various
taxa causing lack of resolution in the strict consensus. In a companion presentation
describing a new species and overall relationships within the subgenus Blackburnia
Sharp (Liebherr and Short 2006), classes of cladograms were excluded because they
defined less tenable biogeographic patterns, or less preferred character-state transfor-
mations. By these means, one preferred cladogram for subgenus Blackburnia was
presented. However this filtering of trees within subgenus Blackburnia left 27 remaining
equally parsimonious topologies for subgenus Metromenus, due to the more ambiguous
character support across that clade (Liebherr and Zimmerman 1998, 2000). For our
purposes, 7 trees among those 27 that supported the overall biogeographic pattern of
progressive colonization of the islands—i.e. the area relationships defined as Kauai
(Oahu (Maui Nui +Hawaii))—were retained for summarization and discussion (Fig. 7).
The inclusion of B. kavanaughi in this cladistic analysis results in a modified
hypothesis of relationships for B. debilis and B. kuiki, as this species pair was previously
placed as adelphotaxon to B. howarthi prior to recognition of B. kavanaughi (e.g.
Liebherr 2003, fig. 8B). Moving B. debilis +B. kuiki to this previous position while
retaining B. kavanaughi as adelphotaxon to the B. epicurus clade, results in a cladogram
one step longer than the shortest 1112-step cladograms (Fig. 7). Moving B. kavanaughi
along with its currently defined adelphotaxon also results in an 1113-step cladogram.
This minimal one-step difference between the two different positions for these taxa
belies the unitary Bremer support values associated with 11 nodes along the spine of the
cladogram network spanning B. huhula to B. howarthi (Fig. 7), demonstrating that
Bremer support cannot be summed over larger sections of a cladogram. Each value is
appropriate only for the node in question, as it merely indicates how much longer a tree
might be than the most parsimonious trees, yet still retain that resolved node (Bremer
The species triplet B. kavanaughi +(B. debilis +B. kuiki) is defined by several
synapomorphies, including longer setae on the antennal pedicel, absence of the anterior
dorsal elytral seta, medially carinate tarsomeres 1–3, and elongate ventrolateral setae on
tarsomere 5. Of these, the absence of the anterior dorsal elytral seta is a uniquely derived
state shared by these three species. Other species within subgenus Blackburnia lack all
three dorsal elytral setae, and two species within subgenus MetromenusB. calathoides
(Sharp) and B. ipu Liebherr—are polymorphic for presence of the hind dorsal elytral
seta, but no other species in the genus matches the setal loss pattern of B. kavanaughi
and allies. Pronotal configuration varies dramatically within this triplet, as the hind
angles of B. kavanaughi are obtuse-rounded (Fig. 1) and those of B. kuiki and B. debilis
angulate (Liebherr and Zimmerman 2000, figs. 192B, C). However, based on how these
characters are coded (Liebherr and Zimmerman 2000) this transformation cannot
countermand the more parsimonious transformations in the above-mentioned
characters, and so we are left to adopt the simplest hypothesis overall (Fig. 7).
Restricting our discussion to extant high Hawaiian Islands, diversification within
subgenus Metromenus unambiguously commenced on the oldest island of Kauai
(Carson and Clague 1995) as evidenced by strong support for B. kahili Liebherr and B.
asquithi as successive adelphotaxa to the remainder of this clade. The position of B.
kavanaughi on the clade subtended by the B. kilauea species quartet (Fig. 7), adds
another isolated representation of Kauai to this portion of the cladogram. Within the B.
kavanaughi triplet, the species occupying the oldest island diverged prior to the other
two species that now occupy fragments of Maui Nui. The isolated position of B.
kavanaughi, among other clades lacking Kauai species, suggests that B. kavanaughi is
the extant representative of a Kauai lineage that colonized a newer island. As the
biogeographic pattern defined by the B. kavanaughi clade fits the general biogeographic
pattern of progressive colonization of newer islands (Liebherr 1997), an undiscovered
member of this clade is predicted for Oahu.
Blackburnia howarthi is a troglobitic inhabitant of lava tube caves in Kipahulu Valley,
Haleakala volcano (Liebherr and Samuelson 1992). Under the newly modified cladistic
hypothesis (Fig. 7), its sister species, B. calathiformis (Sharp), is an epigean species that
ranges across windward Haleakala (Liebherr and Zimmerman 2000, fig. 191). This
revised phylogenetic hypothesis supports most recent divergence of the cave-inhabiting
B. howarthi from sympatric—though topographically overlying—populations of its
epigean adelphotaxon. The timeframe for this event is constrained by the very young
age of the Hana Volcanics, the unit comprising the floor of Kipahulu Valley. The Hana
Unit overlies the Kula Volcanic Unit (Langenheim and Clague 1987), and its flows have
been dated at .0.15–0.12 Ma (Sherrod et al. 2003), with a sample from the Kipahulu
Valley floor dated at 0.121 Ma. The origin of such lava tube caves via differentially
flowing lava ensures their initial sterility. Thus evolutionary divergence of the cave-
adapted B. howarthi has occurred within the last 120,000 yr. Such a pattern represents
the general hypothesis that posits extremely rapid origins and evolution of Hawaiian
cave communities after the lava tubes are emplaced (Howarth 1982, 1987), and requires
any unifying gene flow based on movements of individuals to be overcome by selection
pressures acting on cave populations as they adapt to the newly available subterranean
environments. This restriction is amply supported by the architecture of Hawaiian lava
tube caves, which define extremely long, linear habitats, with only limited or singular
entrances caused by collapse of their ceilings.
Removal of B. debilis and B. kuiki from adelphotaxon status with B. howarthi also
removes a conundrum of speciational timing associated with that prior-hypothesized
relationship. If B. howarthi originated less than 150,000 yr ago as required by geological
evidence, and B. kuiki +B. debilis comprised its adelphotaxon, then divergence of the
latter’s ancestral populations followed by speciation resulting in a Molokai-Haleakala
sister-species pair would need to have occurred since that time. This hypothesis runs
headlong into the establishment of marine incursions that isolated Molokai and Maui
about 0.6 Ma (Price and Elliott-Fisk 2004). Though Molokai was subsequently
connected to Maui via Lanai during more recent interglacial periods, the windward
forest was never directly reconnected. The current placement of B. debilis +B. kuiki as
adelphotaxon to B. kavanaughi places no such unrealistic constraint on speciational
timing or mechanism, as the ancestor of the former two species could have been in place
by 1.2 Ma; the time of maximal development of Maui Nui.
We thank Betsy Harrison Gagne´, State of Hawaii Department of Land and Natural
Resources (DLNR), Natural Areas Reserve System and Alvin Kyono, DLNR, Division of
Forestry and Wildlife, Lihue, Kauai for providing access to natural areas on Kauai. Kokee
Natural History Museum, Hui o Laka—Marsha Erickson director—provided base camp
facilities for Kauai field surveys. I thank Kalani Fronda and Kamehameha Schools for access
to natural habitats on Molokai, and Sam Aruch, The Nature Conservancy of Hawaii
Kamakou Preserve, for his successful rehabilitation of transect 8A from Puu Haha to Uapa.
Dan Polhemus, State of Hawaii DLNR, Division of Aquatic Resources, and Curtis Ewing
and Andrew Short, Cornell University collaborated in the field. Dan Otte provided his
distributional map blank. National Science Foundation REVSYS award DEB-0315504
supported this research.
Bremer, K. 1994. Branch support and tree stability. Cladistics 10: 295–304.
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a Hot Spot Archipelago. Smithsonian Institution Press, Washington.
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Coleoptera). Pacific Science 51: 424–439.
Liebherr, J. K. 2001. Blackburnia gastrellariformis sp. n. (Coleoptera: Carabidae), from Molokai:
successful prediction of a new taxon by reconciled tree analysis. Insect Systematics and
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Liebherr, J. K. 2003. Blackburnia lata sp. n. (Coleoptera: Carabidae) from Kauai: morphological
transformation in the arboreal microhabitat. Insect Systematics and Evolution 34: 41–52.
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Hawaiian lava tubes (Coleoptera: Carabidae). Pan-Pacific Entomologist 68: 157–168.
Liebherr, J. K. and A. E. Z. Short. 2006. Blackburnia riparia, new species (Coleoptera: Carabidae,
Platynini): a novel element in the Hawaiian riparian insect fauna. Journal of the New York
Entomological Society 114: 1–16.
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Hawaiian Platynini (Coleoptera: Carabidae). Systematic Entomology 23: 137–172.
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Introduction and Tribe Platynini. Insects of Hawaii 16: 494 pp.
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¯, East Maui, a post-shield-stage volcano of
the Hawaiian island chain. Geological Society of America Bulletin 115: 683–694.
Received 1 May 2006; accepted 12 July 2006.
... The biogeographic histories of Hawaiian and Society island taxa in the carabid tribes Platynini and Moriomorphini also involve direct colonisation from the south-west Pacific without any stepping-stone relationships. The former comprises species of the Hawaiian endemic platynine genus Blackburnia Sharp (Liebherr and Zimmerman 2000;Liebherr 2001Liebherr , 2003Liebherr , 2007Liebherr and Short 2007) versus the Tahitian platynine fauna comprising only three species (Perrault 1977). Hawaiian colonisation by Blackburnia is hypothesised to have occurred 28-30 Mya, long before the origin of Kauai at 5.1 Mya, based on the lineage's phylogenetic relationships to other Australian and Pacific taxa (Liebherr 2005). ...
Bembidion (Sloanephila) tahitiense, sp. nov. is described from Mont Mauru, an isolated massif of Tahiti Nui volcano. Based on evidence from seven genes (four nuclear protein-coding, one mitochondrial protein-coding, two nuclear ribosomal), its sister group is the Australian B. jacksoniense Guerin-Meneville, with which it shares a synapomorphic spur on the ostium of the male genitalia. In contrast to B. jacksoniense, B. tahitiense is brachypterous, with rounded humeri, constricted posterior pronotal margins and convex body form. Examination of the seven genes in two species of the Hawaiian subgenus Nesocidium Sharp reveals that the sister group of Nesocidium is subgenus Zecillenus Lindroth from New Zealand. These two subgenera belong to the Ananotaphus complex, a clade inhabiting Australia, New Zealand and Hawaii. The relationships of the second Hawaiian subgenus, Gnatholymnaeum Sharp, are less clear, although Gnatholymnaeum belongs to the Bembidion series (along with Sloanephila and the Ananotaphus complex). Bembidion beetles colonised the Society and Hawaiian islands independently from source areas in the south-west Pacific. Based on parsimonious reconstructions of flight-wing configuration, the Tahitian and Hawaiian colonisations involved winged individuals. Colonisation of the Society and Hawaiian islands by carabid beetles of two other tribes - Platynini and Moriomorphini - follow the dispersal patterns hypothesised for Bembidion.
... The newly described species of Blackburnia were analysed cladistically by adding them to the previously developed cladistic dataset based on Liebherr and Zimmerman (1998), that matrix subsequently modified by Liebherr (2001Liebherr ( , 2003Liebherr ( , 2006 and Liebherr and Short (2006). The original matrix included information based on 206 characters, with those characters described in Liebherr and Zimmerman (1998). ...
Conference Paper
The Hawaiian Islands are home to over 5,000 described, indigenous insect species. Intense phylogenesis has resulted in species radiations that serve as models for evolutionary biology. The genus Blackburnia (Coleoptera: Carabidae) has been the subject of numerical cladistic analysis characterized by 1, inclusion of all known species, 2, comprehensive morphological character selection; and 3, a test for monophyly using multiple outgroups. Phylogenetic analysis of extant species can now be complemented via the characterization of a pre-Polynesian insect community recovered from Makauwahi Cave, south-coastal Kauai. This community included both subsequently extirpated and currently extant carabid beetle taxa. Inclusion of sub-fossil taxa in the cladistic analysis of Blackburnia improves phylogenetic resolution while leaving previous taxic relationships undisturbed. Phylogenetic placements of sub-fossil taxa are uniformly in accord with area relationships derived from items-of-error biogeographic analysis based on extant species. Presence of sub-fossils representing extant species in the Makauwahi Cave deposit allows an assessment of the changes to biotic distributions that have occurred during the history of Kauais human habitation.
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Three major eruptive stages in a Hawaiian volcano's life-shield stage (tholeiitic), postshield stage (alkalic), and rejuvenated stage (alkalic)-have generally provided a basis for dividing the volcanic rocks into stratigraphic units. The stratigraphic nomenclature of the Hawaiian Islands is herein reviewed and updated. We give in tabular form a brief summary of each stratigraphic unit, including its lithology, occurrence, thickness, type and reference localities, stratigraphic relations, age, and any stratigraphic changes made herein.-from Authors
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Blackburnia riparia, new species is described from the summit of Mt. Waialeale, Kauai, Hawaii, incorporated into a phylogenetic analysis of Blackburnia Sharp, and thereby placed as the adelphotaxon to two other allopatric Kauai species, B. lata Liebherr and B. atra Liebherr. The new species occurs in riparian habitats, including vertical rock-faced seeps and algal mats, and is documented to feed on aquatic larvae of Micropsectra Kieffer (Diptera: Chironomidae). Blackburnia elegans (Sharp) adults also occur in streams on Waialeale summit where they were found walking under water on the undersides of large, flat rocks. Based on documented prey and observed behavior, both species appear to forage under the waterline. The abdominal and elytral anatomy of adult B. riparia and B. elegans is suitable for retention of a subelytral air bubble, suggesting these species respire underwater. Adults and associated larvae of Blackburnia mandibularis Liebherr inhabit moss-mats along falls' margins at Waipoo Falls, Kokee State Park, Kauai, indicating that this species' active life stages also reside in the riparian zone. Attributes of the larval head capsule and mandibles for B. riparia and B. mandibularis are associated with homologous characters expressed in adult anatomy, ontogenetically linking larval and adult head and mouthpart specializations. The various evolutionary origins of riparian habits are examined across Blackburnia. Via one route, occupation of the montane riparian zone has evolved from terrestrial moss-mat habitation, with concomitant evolutionary reduction of adult compound eyes. A second evolutionary route to the riparian zone is based on occupation of open, disturbed or ephemeral habitats, in some cases assisted by adult winged dispersal.
The carabid beetle species of the tribe Platynini are revised for the Hawaiian Islands based on information from the adult stage. The 129 known species are considered member taxa of the genus Blackburnia Sharp, NEW STATUS . Twenty-two generic names previously used to classify the Hawaiian Platynini are placed in synonymy with Blackburnia. Primary types representing all available species and varietal names were examined, and lectotypes designated. Native species are arrayed in four subgenera: Protocaccus, NEW SUBGENUS, comprising one species, B. (P.) mandibularis, NEW SPECIES, of Kauai; Colpocaccus Sharp, NEW STATUS, with four species; Blackburnia, sensu stricto, with 51 species, of which 13 are newly described; and Metromenus Sharp, NEW STATUS, with 73 species, of which 19 are newly described. NEW SPECIES of the subgenus Blackburnia include B. atra, B. bryophila, and B. waialeale of Kauai; B. hihia of Oahu; B. polhemusi of Molokai; B. viridis, B. pukalaina, B. pilikua, B. polipoli, B. medeirosi, and B. kipahulu of Maui; and B. kukui and B. ewingi of Hawaii Island. NEW SPECIES of the subgenus Metromenus include B. kahili, B. asquithi, and B. pauma of Kauai; B. huhula, B. paludicola, B. paloloensis, and B.fordi of Oahu; B. platyophthalmica and B. hakeakapa of Molokai; B. ulaula, B. lihau, B. ipu, B. kuiki, B. abaxoides, B. auana, B. foveolata, B. bartletti, B. kauwa, and B. fulgida of Maui. NEW NAMES required to replace homonyms resulting from generic synonymization include: Blackburnia (Blackburnia) kamehameha = AnchotefJlus gracilis Sharp, 1903; Blackburnia (Metromenus) komohana = Mecomenus koebelei Sharp, 1903; Blackburnia (Metromenus) haleakala = Mecostomus perkinsi Sharp, 1903; Blackburnia (Metromenus) kilauea = Atelothrus hawaiiensis Sharp, 1903; Blackburnia (Metromenus) lanaihalensis = Mesothriscus lanaiensis Sharp, 1903. The following NEW SYNONYMIES are proposed (senior synonyms first): Blackburnia (Colpocaccus) posticata (Sharp) = Colpocaccus marginatus Sharp; Blackburnia (Colpocaccus) hawaiiensis (Sharp) = Colpocaccus apicalis Sharp; Blackburnia (Blackburnia) lucipetens (Blackburn) = Colpodiscus lahainensis Sharp; Blackburnia (Blackburnia) micans (Sharp) = Derobroscus politus Sharp and Derobroscus solitarius Sharp; Blackburnia (Blackburnia) brevipes (Sharp) = Disenochus flavitarsis Sharp; Blackburnia (Metromenus) muscicola (Blackburn) = Mesothriscus prognathus Sharp; Blackburnia (Metromenus) proterva (Blackburn) = Metromenus angustifrons Sharp; Blackburnia (Metromenus) kilauea =Mesothriscus hawaiiensis Sharp (junior homonym of Colpocaccus hawaiiensis Sharp) and Metromenus lentus Sharp (junior homonym of Pseudobroscus lentus Sharp); Blackburnia (Metromenus) fraterna Blackburn = Anchomenus scrupulosus Blackburn, Metromenus aequalis Sharp, and Metromenus velox Sharp; Blackburnia (Metromenus) pavida Sharp = Mesothriscus truncatus Sharp; Blackburnia (Metromenus) sphodriformis (Sharp) = Atelothrus limbatus Sharp, Atelothrus longicol/is Sharp, Mesothriscus collaris Sharp, and Metromenus cinctus Sharp; Blackburnia (Metromenus) erro (Blackburn) = Atelothrus politus Sharp; Blackburnia (Metromenus) insociabilis (Blackburn) = Atelothrus stenopus Sharp. Identification keys are provided to the tribes of Carabidae known from Hawaii, subgenera and species of native Hawaiian Platynini, and platynine species introduced to Hawaii. Native species treatments include diagnosis and description for newly recognized species or diagnostic combination for previously proposed species, line illustrations and habitus photo, distributional map, and natural history and phylogenetic information. Taxonomic classification of the Hawaiian Platynini is based on cladistic analysis. The evolution ofbrachyptery has proceeded four times independently during the course of the platynine radiation. Two species are known cave troglobites, exhibiting reduced eyes and reduced cuticular melanization. These species are most closely related to moss-inhabiting taxa, suggesting that colonization of moss-covered voids in the forest floor was the means to enter lava tube caves. Phylogenetic relationships among the native species indicate that Kauai is the root of the current extant radiation, with progressive colonization of the Hawaiian Islands continuing throughout time since the origin of Kauai. Within-island vicariance has facilitated speciation on all of the islands. On the older island of Oahu, this has been followed by secondary dispersal, resulting in many broadly sympatric species within the Waianae and Koolau Ranges. This secondary mixing has been largely restricted to present rain forest habitats, as 22 of the 32 Oahu species remain restricted to either the Waianaes or the Koolaus. Vicariant barriers are especially evident on the sunken and fragmented island of Maui Nui, where endemic relatives can often be found on Molokai, West Maui, and Haleakala. Haleakala is the site of the most loss of biodiversity, through the loss or disturbance of mesic forests of koa (Acacia koa A. Gray) that housed numerous species collected in the 1870s to 1890s. On all the islands, evident losses of species are concentrated in the drier, leeward communities. The most dramatic reduction in populations has been the loss of B. (Colpocaccus) tantalus from Oahu. Though this species was distributed across the island, and constituted 40% of specimens in the nineteenth century, it has not been seen since 1940. Three adventive platynine species are also treated taxonomically: Laemostenus complanatus (Dejean), Calathus rujicollis (Dejean), and Metacolpodes buchanani (Hope). The first colonized Hawaii Island about 1950 and has expanded its range throughout much of the drier forests of Mauna Kea and Mauna Loa. Calathus ruficollis, a native of the Pacific coast of North America, is found on Oahu, and was probably introduced from San Diego. Metacolpodes buchanani, native to Asia but also introduced to Oregon, was introduced to Oahu in or before 1991, and by 1999 had expanded its range to include Kauai, Molokai, Maui, and the island of Hawaii.
Characters of the female reproductive tract, ovipositor, and abdomen are analyzed using cladistic parsimony for a comprehensive representation of carabid beetle tribes. The resulting cIadogram is rooted at the family Trachypachidae. No characters of the female reproductive tract define the Carabidae as monophyletic. The Carabidae exhibit a fundamental dichotomy, with the isochaete tribes Metriini and Paussini forming the adelphotaxon to the Anisochaeta, which includes Gehringiini and Rhysodini, along with the other groups considered member taxa in Jeannel's classification. Monophyly of Isochaeta is supported by the groundplan presence of a securiform helminthoid sclerite at the spermathecal base, and a rod-like, elongate laterotergite IX leading to the explosion chamber of the pygidial defense glands. Monophyly of the Anisochaeta is supported by the derived division of gonocoxa IX into a basal and apical portion. Within Anisochaeta, the evolution of a secondary spermatheca-2, and loss of the primary spermatheca-I has occurred in one lineage including the Gehringiini, Notiokasiini, Elaphrini, Nebriini, Opisthiini, Notiophilini, and Omophronini. This evolutionary replacement is demonstrated by the possession of both spermatheca-like structures in Gehringia o/ympica Darlington and Omophron variegatum (Olivier). The adelphotaxon to this spermatheca-2 clade comprises a basal rhysodine grade consisting of Clivinini, Promecognathini, Amarotypini, Apotomini, Melaenini, Cymbionotini, and Rhysodini. The Rhysodini and Clivinini both exhibit a highly modified laterotergite IX; long and thin, with or without a clavate lateral region. This may represent a synapomorphous derivation, or convergence based on a tubular abdomen and burrowing habit. The basal grade gives rise to a grade of taxa sharing the presence of a ramus--a sclerotized portion of the vaginal wall situated medially to each gonocoxal base--along with the plesiomorphic spermatheca-I. The previous interpretation of the gonocoxal rami as a groundplan feature of Carabidae representing portions of abdominal segment VIII is rejected based on its derived occurrence relative to the origin of the family. The gonocoxal ramus or cicindine grade includes I) Cicindini, 2) a monophyletic "carabine" clade including Siagonini, Cychrini, Pamborini, Carabini, and Cicindelini, 3) and the Scaritini and Hiletini which are paraphyletically related to a monophyletic group exhibiting the harpalidian abdominal configuration, including Broscini and all remaining tribes of Carabidae. Broscini retain the separate accessory gland and spermathecal arrangment of Scaritini and Hiletini, and comprise the adelphotaxon to the other remaining tribes. Patrobini represent the next divergent group, exhibiting an appended spermathecal gland but retaining the ramus at the base of gonocoxite I. The remaining tribes, including member tribes of Jeannel's Stylifera (in part) and Conchifera, are ambiguously related based on female characters at a highly polytomous II node. Within this polytomous clade, Pseudomorphini share a derived basal spermathecal sclerite with Geobaenini, Lachnophorini, and Odacanthini. Cnemalobini are placed as the adelphotaxon to Morionini. Other tribal relationships remain ambiguous due to basic homogeneity of the female reproductive tract observed throughout these tribes. The problematic Brachinini appear best placed among these tribes, although a less preferred but equally parsimonious placement is as adelphotaxon to Clivinini. Psydrini and Zolini appear polyphyletic based on female characters, with constituent subtribes placed at various positions between the gonocoxal ramus grade and the highest polytomous clade. The basal position of Clivinini and Apotomini supported by female characters is contrasted with Jeannel's placement in Scrobifera and Stylifera, respectively, based on thoracic structure.
The Parsimony Ratchet1 is presented as a new method for analysis of large data sets. The method can be easily implemented with existing phylogenetic software by generating batch command files. Such an approach has been implemented in the programs DADA (Nixon, 1998) and Winclada (Nixon, 1999). The Parsimony Ratchet has also been implemented in the most recent versions of NONA (Goloboff, 1998). These implementations of the ratchet use the following steps: (1) Generate a starting tree (e.g., a “Wagner” tree followed by some level of branch swapping or not). (2) Randomly select a subset of characters, each of which is given additional weight (e.g., add 1 to the weight of each selected character). (3) Perform branch swapping (e.g., “branch-breaking” or TBR) on the current tree using the reweighted matrix, keeping only one (or few) tree. (4) Set all weights for the characters to the “original” weights (typically, equal weights). (5) Perform branch swapping (e.g., branch-breaking or TBR) on the current tree (from step 3) holding one (or few) tree. (6) Return to step 2. Steps 2–6 are considered to be one iteration, and typically, 50–200 or more iterations are performed. The number of characters to be sampled for reweighting in step 2 is determined by the user; I have found that between 5 and 25% of the characters provide good results in most cases. The performance of the ratchet for large data sets is outstanding, and the results of analyses of the 500 taxon seed plant rbcL data set (Chase et al., 1993) are presented here. A separate analysis of a three-gene data set for 567 taxa will be presented elsewhere (Soltis et al., in preparation) demonstrating the same extraordinary power. With the 500-taxon data set, shortest trees are typically found within 22 h (four runs of 200 iterations) on a 200-MHz Pentium Pro. These analyses indicate efficiency increases of 20×–80× over “traditional methods” such as varying taxon order randomly and holding few trees, followed by more complete analyses of the best trees found, and thousands of times faster than nonstrategic searches with PAUP. Because the ratchet samples many tree islands with fewer trees from each island, it provides much more accurate estimates of the “true” consensus than collecting many trees from few islands. With the ratchet, Goloboff's NONA, and existing computer hardware, data sets that were previously intractable or required months or years of analysis with PAUP* can now be adequately analyzed in a few hours or days.
The postshield and previously inferred rejuvenated-stage history of Haleakalā volcano is reevaluated on the basis of 52 new K-Ar ages, 42 from the postshield Kula Volcanics and 10 from the overlying Hāna Volcanics. Postshield extrusion was robust from 0.93 to 0.76 Ma. A period of low extrusion rate or volcanic quiescence occurred between 0.76 and 0.65 Ma, well within Kula time. A chemical change to increasingly alkalic lava occurred at this time as the volcano changed from broadly hawaiitic to basanitic in its eruptive products and robust extrusion resumed. A slightly longer period of low extrusion rate or quiescence occurred after ca. 0.4 Ma, but only trifling change in geochemical character is observed. Geochemically, the Hāna Volcanics unit, chiefly basanitic, overlaps greatly with the upper part of the Kula Volcanics; there is a weak tendency to slightly more alkaline character among the Hāna Volcanics. The age of the Kula/Hāna boundary is ca. 0.15-0.12 Ma; thus, volcanic quiescence of only ∼0.03 m.y. separates the two formations, much shorter than the previously known limit of 0.25-0.30 m.y. The brevity of this hiatus, coupled with coincident vent loci and broadly similar geochemical characteristics for the Hāna and the upper part of the Kula Volcanics, indicates that the Hāna Volcanics unit comprises deposits of postshield-stage volcanism that has waned substantially since ca. 0.4-0.3 Ma. Haleakalā has not yet begun a classically defined rejuvenated stage. Our findings support recent numerical modeling of plume-lithosphere interactions that predict that Haleakalā is near the end of its postshield growth.
Specimens taxonomically treated in the Fauna Hawaiiensis were associated by cluster analysis, thereby reconstructing assemblages of Hawaiian carabid beetle species (Coleoptera: Carabidae) observed during the late 19th century. Associations among specimens representing 193 species permit concise hypotheses of habitat preferences for many of the 32 carabid species collected during the early period of European scientific exploration (1872–1902), but not observed since. These associations are consistent with data derived from contemporary biological surveys of Hawai'i. Absence of entire clusters of associated species from recent collections suggests actions of common agents leading to extinction or extreme population reduction. The candidate list of threatened and endangered species of the US. Fish and Wildlife Service established prior to 1994 included one Hawaiian carabid species missing since 1902, versus eight other species collected at various times over the past century. Improvements in knowledge of carabid beetle species’ spatial distribution and temporal persistence derived from recent field survey and taxonomic research demonstrate that the types of criteria used to construct that list must be rejected. Future consideration of official conservation status for any Hawaiian carabid beetle species must take into account the status of ecologically associated species, and the limited likelihood that individuals of all extant species can be consistently observed in nature due to their natural relative rarity or their secretive habits within restricted geographic and ecological distributions. Historical specimen associations serve as the best guides for continuing efforts to monitor known faunal members and to rediscover long‐missing species. These associations also serve to link information concerning individual species with particular habitats.
Blackburnia lata sp. n. is described from the western reaches of the Alakai Swamp Trail, Kauaikinana Stream watershed, Kauai. Blackburnia lata is placed phylogenetically among other Kauai species in the subgenus Blackburnia Sharp, 1878. It is sympatric with three closely related species, all distributed largely within the limits of the primeval Kauai caldera, but allopatric with its adelphotaxon, B. atra Liebherr, found on Mt. Kahili, an isolated southern spur of the Mt. Waialeale massif. All five species occur in arboreal mossmats, at the same time exhibiting substantial variation in tarsal configuration. Tarsal evolution across this mossmat-specific lineage suggests that a single tarsal conformation is not required for occupation of this arboreal ecological zone. Historical transformation patterns of tarsal characters are not entirely congruent with the phylogenetic relationships among these species, and inclusion or exclusion of these characters in cladistic analysis is irrelevant to the definition of phylogenetic relationships. This incongruence argues against the necessity of excluding characters from phylogenetic analysis because they are deemed adaptive, while supporting the total evidence approach, wherein all potential synapomorphies are analyzed simultaneously to estimate phylogeny, regardless of their putative adaptive significance.