El Genero Creophilus

Zoological Journal of the Linnean Society, 2011, 163, 723–812.
With 37 figures
Testing the phylogenetic utility of morphological

character systems, with a revision of Creophilus Leach
(Coleoptera: Staphylinidae) zoj_725 723..812
DAVE J. CLARKE1,2*
1
Department of Zoology, The Field Museum of Natural History, 1400 S Lake Shore Drive, Chicago,
IL 60605, USA
2
Department of Biological Sciences, University of Illinois, at Chicago, 845 West Taylor St. M/C 066,
Chicago, IL 60607, USA
Received 29 July 2010; revised 16 November 2010; accepted for publication 6 December 2010
Many studies have examined the phylogenetic utility of different kinds of molecular data, and have often compared
these with morphology. However, relatively few studies have phylogenetically evaluated different morphological
character systems. Using the Creophilus complex, the phylogenetic utility of external structural characters, male
genitalia, female genitalia, and chaetotaxy of adults is examined for the first time in the megadiverse beetle family
Staphylinidae. A data set of 121 phylogenetically informative characters of 24 terminal taxa was analysed
separately and simultaneously using parsimony and the phylogenetic utility of each partition was compared with
widely used statistics and support measures. External structures had the least homoplasy, resolved the most nodes
separately, and largely determined the topology of the simultaneous analysis. Male and female genitalia contrib-
uted phylogenetic signal mostly congruent with external characters. Despite extensive homoplasy, chaetotaxy
contributed the majority of hidden support, and was critical for resolving several terminal nodes in the simulta-
neous analysis. All character systems were informative throughout the tree, and when combined provided the
best-supported hypothesis. However, there is a need to distinguish hidden support from dispersion of homoplasy
when combining data sets of varying quality. The Creophilus complex, Creophilus Leach, Liusus Sharp, and the
Creophilus erythrocephalus (Fabricius) and C. maxillosus (Linnaeus) species-groups are each strongly supported
monophyletic groups. Hadrotes Mäklin is not monophyletic and Hadrotes wakefieldi Cameron is the sister-species
of the rest of the Creophilus complex, but included within it. Creophilus is revised to include 12 species, including
Creophilus galapagensis sp. nov. and C. rekohuensis sp. nov. Creophilus huttoni (Broun) is removed from
synonymy with C. oculatus (Fabricius), and C. insularis (Fauvel), C. villipennis Kraatz, and C. violaceus (Fauvel)
are synonymized under C. flavipennis (Hope) comb. et stat. nov.
© 2011 The Linnean Society of London, Zoological Journal of the Linnean Society, 2011, 163, 723–812.
doi: 10.1111/j.1096-3642.2011.00725.x
ADDITIONAL KEYWORDS: Bremer support – internal sac – macroseta – necrophilous – rove beetle –
Staphylinina – Staphylininae – Staphylinini – taxonomic revision.
INTRODUCTION absence of such data, evolutionary hypotheses and

systematic studies must rely on data from other
For the vast majority of species, molecular data will
sources. Although morphology continues to contribute
not be available in the near futrue, or indeed ever
positively to systematics, especially in entomology
(Novacek & Wheeler, 1992; Laurance, 2007). In the
(Wheeler, 2008; Bybee et al., 2010), the role it should
play is persistently controversial (Scotland, Olmstead
& Bennett, 2003; Tautz et al., 2003; Jenner, 2004;
*E-mail: [email protected] Smith & Turner, 2005; Will, Mishler & Wheeler,
© 2011 The Linnean Society of London, Zoological Journal of the Linnean Society, 2011, 163, 723–812 723
724 D. J. CLARKE
2005). Insect systematics, however, still relies largely chaetotaxy. Most studies lack discussion of why
on morphology for phylogenetic inference and a large various character systems were included or excluded
majority of endopterygote insect phylogenies are (but see Solodovnikov, 2006 for an example of why
based solely on morphology, and most of these on the chaetotaxy was excluded, and Frisch, Burckhardt &
morphology of adults (Meier & Lim, 2009). In general, Wolters, 2002 for a surprising example of why few
insect organismal data can comprise characters of the structural characters were included), or the degree of
exoskeleton and its various cuticular structures (e.g. existing variation not utilized in different character
Harris, 1979; Richards & Richards, 1979; Solodovni- systems. Hence it is increasingly difficult to judge the
kov, 2007), wings and appendages (e.g. Kukalová- quality and extent of phylogenetically informative
Peck & Lawrence, 2004), internal structures and variation in various groups. In molecular systematics
anatomy (e.g. Beutel & Vilhelmsen, 2006; Friedrich & there has been a great deal of attention devoted to
Beutel, 2006), behaviour (e.g. Wenzel, 1992), and assessing the relative phylogenetic utility of morpho-
ecology (e.g. Miller & Wenzel, 1995). Like DNA, some logical versus molecular characters (e.g. Baker,
of these apply to all semaphoronts and therefore can Xiaobo & Desalle, 1998; Wahlberg et al., 2005;
provide a huge array of phylogenetic information. Wortley & Scotland, 2006) and the relative phyloge-
Adults can provide a rich variety of structures for use netic utility of different genes for phylogenetic infer-
in both taxonomy and phylogeny in many groups, and ence (e.g. Simon et al., 1994; Baker, Wilkinson &
the importance of male genitalia in insect taxonomy DeSalle, 2001; Wild & Maddison, 2008). This trend
is widely known (Eberhard, 1985). In addition, has not yet been mirrored so extensively in phyloge-
insect phylogenetic analyses sometimes incorporate netic studies based on non-molecular traits, and there
a large number of male genital characters, conferring are comparatively fewer papers quantitatively testing
a very broad significance to this organ system in the phylogenetic utility of alternative character
systematics. systems. Most entomological studies have evaluated
In the enormous beetle family Staphylinidae, with adult and immature characters, and no comparison of
more than 55 000 described species and growing the various adult character systems has yet been
rapidly (Newton, 2007; Grebennikov & Newton, performed in Staphylinidae, a group for which the
2009), morphology still dominates phylogenetic majority of phylogenies are based on adult morphol-
inference. Adult characters dominate both staphylinid ogy alone. Although quality control is a critical issue
taxonomy and phylogenetics, in part because of the (Rieppel & Kearney, 2002), if the maximum value of
gaping chasm of knowledge between adult and morphological characters is to be realized in phyloge-
larval systematics (Newton, 1990, 2007). This reflects netic inference, it is clear that existing character
broader trends of under-utilized immature character systems need to be more fully utilized, previous
systems in endopterygote systematics (Meier & Lim, homology statements refined and reanalysed (Jenner,
2009), and the nearly complete lack of other organis- 2004), and ‘unconventional’ character sources more
mal data such as behaviour (but see Danoff-Burg, extensively explored and tested. Smith & Turner
2002). The literature on rove beetle phylogenetics (2005) and others argue convincingly against claims
(discussed in detail later) indicates that, specifically, that morphology has limited value for phylogeny
external cuticular structures have contributed the reconstruction in the ‘molecular age’ (Scotland et al.,
most characters to adult morphological analyses 2003) and highlight that indeed most of the relevant
whereas many other character systems seem gener- morphological variation still remains untapped.
ally poorly represented or entirely omitted. This lit- For the first part of this study, the major goal was
erature indicates: (1) relatively small character/taxon to demonstrate the diversity of undiscovered charac-
ratios in most studies; (2) a general under-utilization ters available even in a comparatively well-known
of many character systems, especially female genita- group of rove beetles, and to compare the relative
lia and structures associated with the male internal phylogenetic utility of characters taken from tradi-
sac, some being completely or nearly completely unex- tionally delimited morphological character systems. A
plored (e.g. internal anatomy); (3) variability and phylogenetic analysis of the species belonging to the
inconsistencies in patterns of character usage (espe- Creophilus complex is used here in the first analysis
cially chaetotaxy) relating to taxonomic group; and (4) of the phylogenetic utility of male genitalia, female
discrepancies between the use of adult and larval genitalia, and adult chaetotaxy in relation to external
chaetotaxy when both semaphoronts are analysed. structural characters in rove beetles. This compara-
The reasons for these patterns remain unclear and tively small group comprises several morphologically
ultimately it is difficult to conclude anything from the closely related genera, exhibits a diverse range of
literature regarding the absolute and relative phylo- morphological characters, and is therefore an ideal
genetic utility of different character systems, espe- group to pursue such a test. The following questions
cially male genitalia, female genitalia, and adult are addressed. (1) Do male genitalia, female genitalia,
© 2011 The Linnean Society of London, Zoological Journal of the Linnean Society, 2011, 163, 723–812
PHYLOGENETIC UTILITY OF CHARACTERS IN CREOPHILUS 725
and chaetotaxy contribute more noise than signal to a fauna (Levot, 2003; Archer, 2004; Nowak, 2004), and
simultaneous analysis of all morphological data (con- historically has promoted interest in using these
sistent with the concept of ‘holomorphology’: Hennig, beetles for biocontrol purposes (Fullaway, 1923;
1966)? (2) According to various measures of phyloge- Swezey, 1923). Most historical or contemporary treat-
netic utility, how do these character systems compare ments and identification keys omit more than half the
with external structural characters and among them- species or are based only on C. maxillosus (e.g. Moore
selves? (3) Are there any patterns regarding the dis- & Legner, 1979; Naomi, 1982; Smetana & Davies,
tribution of support, i.e. do certain character systems 2000), and none has included all austral species.
provide more support to some nodes than to others? There are still several long-standing taxonomic prob-
To address these questions I apply methods developed lems, and given the size and prominence of these
by several authors (e.g. Gatesy, O’Grady & Baker, beetles, the genus is a compelling example of a sta-
1999) to explore the phylogenetic utility of alternative phylinid group in need of a modern synthesis. It
character systems, and to examine and explain is also a sobering illustration of the amount of
hidden support from different data sets that may basic taxonomic work still needed in Staphylinidae.
emerge in a combined analysis of all data. I also Improved taxonomic and biological knowledge of
compare the data set developed here to a broad these species is therefore broadly useful, especially as
sample of rove beetle morphology-based phylogenetic several species occur sympatrically, look very similar,
analyses. and are of interest in applied entomology. In a revi-
sion of Creophilus, the second part of this study
reinforces the taxonomic value of male genitalia
THE CREOPHILUS COMPLEX AND THE GENUS through detailed study of the internal sac, and also
CREOPHILUS LEACH, 1819 demonstrates the comparable taxonomic value of
The ‘Creophilus complex’, as defined here, includes female genitalia and chaetotaxy in Creophilus.
Creophilus (12 spp.), Liusus Sharp, 1889 (2 spp.),
Hadrotes Mäklin, 1852 (2 spp.), and the monotypic
genera Hadropinus Sharp, 1889 and Thinopinus
MATERIAL AND METHODS
LeConte, 1852 (henceforth, Hadrotes species are
abbreviated to Ht. crassus and Ht. wakefieldi, and COLLECTION ABBREVIATIONS AND MATERIAL
Hadropinus fossor Sharp, 1889 to Hp. fossor). Eight of EXAMINED
the 18 species in this group occur in south temperate Collection abbreviations are taken mostly from
or Gondwanan regions, with the rest distributed vari- Arnett, Samuelson & Nishida (1993) and are given in
ously in northern hemisphere or tropical regions. capitals. Additional collections not listed there are
Eleven of the 12 Creophilus species and Hadrotes taken from Naumann, Jusoh & Lumb (2003), and
wakefieldi Cameron, 1945 are fully terrestrial, the acronyms for private collections are given in upper
others exclusively intertidal. The systematic goals and lower case. Listed here are the institutions from
of the analysis were therefore to test the monophyly which I borrowed specimens, plus other institutions
of (1) the Creophilus-complex, (2) the genera mentioned in the text (marked with an asterisk):
Creophilus, Hadrotes, and Liusus, (3) the informal AMNH (American Museum of Natural History,
erythrocephalus- and maxillosus-groups within Creo- New York City, USA); AMNZ (Auckland Institute &
philus (Newton, 1985), and (4) the beach-dwelling Museum, Auckland City, New Zealand); AMS (Aus-
species in Hadrotes, Hadropinus, Liusus, and tralian Museum, Sydney, Australia); ANIC (Austra-
Thinopinus. lian National Insect Collection, Canberra, Australia);
The type species of Creophilus Leach 1819, C. max- BMNH (The Natural History Museum, London,
illosus (Linnaeus, 1758), was described more than 250 UK); BPBM (Bernice P. Bishop Museum, Honolulu,
years ago in the Systema naturae, and has since been USA); CAS (California Academy of Sciences,
widely studied taxonomically, morphologically, and San Francisco, USA); CMNC (Canadian Museum
ecologically (see bibliography in Herman, 2001b). The of Nature, Ottawa, Canada); CMNZ (Canterbury
12 species now included in Creophilus are some of Museum, Christchurch, New Zealand); CNC (Cana-
the largest (up to c. 30 mm) and more attractively dian National Collection of Insects, Ottawa, Canada);
coloured rove beetles commonly encountered (Fig. 1). DEI (Senckenberg Deutsches Entomologisches Insti-
Adults and larvae of nearly all species are primarily tut, Müncheberg, Germany); FMNH (Field Museum
predators of maggots and are found almost exclu- of Natural History, Chicago, USA); IRSNB (Institut
sively at carrion. The biology and ecology of these Royal des Sciences Naturelles de Belgique, Brussels,
beetles has attracted attention from forensic ento- Belgium); CDRS (Charles Darwin Research Station,
mologists concerned with estimating minimum post- Galapagos Islands, Ecuador); LUNZ (Lincoln Univer-
mortem intervals using succession of necrophilous sity Entomological Research Museum, Lincoln, New
726 D. J. CLARKE
Figure 1. Habitus photographs of selected species in the Creophilus complex (males, except as indicated). A, C.
erythrocephalus; B, C. huttoni; C, C. oculatus; D, C. rekohuensis; E, C. albertisi; F, C. maxillosus maxillosus; G, C. incanus;
H, C. galapagensis; I, C. flavipennis (Indonesia: Sulawesi); J, C. variegatus, female; K, Hadropinus fossor; L, Hadrotes
crassus, female; M, Thinopinus pictus; N, Liusus hilleri; O, Hadrotes wakefieldi. Scale bars = 5 mm.
䉳
Zealand); MAMU (Macleay Museum, University of villosus (Gravenhorst, 1802) and some other species
Sydney, Sydney, Australia); MBBJ* (Museum Zoologi- accumulated and determined by A. F. Newton
cum Bogoriense, Bogor, Indonesia); MCSN (Museo (FMNH) (most not examined by me). Material exam-
Civico di Storia Naturale ‘Giacomo Doria’, Genoa, ined data and a complete institutional list for all
Italy); MCZ (Museum of Comparative Zoology, material that this study is based on are available as
Harvard University, Cambridge, MA, USA); MHNG supporting information, Appendix S1.
(Muséum d’Histoire Naturelle, Geneva, Switzerland);
MNNC (Museo Nacional de Historia Natural, San- SPECIMEN PREPARATION AND EXAMINATION
tiago, Chile); MONZ (Museum of New Zealand – Te Both dried and alcohol-preserved specimens were
Papa Tongarewa, Wellington, New Zealand); MMUE examined with a dissecting microscope (Leica
(Manchester Museum, The University, Manchester, MZ12.5), and all line illustrations were created with
UK); MZHF (Finnish Museum of Natural History, the aid of a camera lucida attachment. Specimens
Helsinki, Finland); NMW (Naturhistorisches Museum were gold-coated for scanning electron microscopy
Wien, Vienna, Austria); Nunn (John Nunn private (SEM) and examined with an LEO EVO 60 environ-
collection, Dunedin, New Zealand); NZAC (New mental scanning electron microscope. Habitus and
Zealand Arthropod Collection, Landcare Research, other photographs were taken with a Microptics ML
Auckland); OMNZ (Otago Museum, Dunedin, New Macro XLT digital imaging system with several
Zealand); OSAC* (Oregon State Arthropod Collection, images amalgamated using Helicon Focus. Wing
Oregon State University, Corvallis, OR, USA); QCAZ* preparations were made by dissecting the left wing at
(Quito Catholic Zoology Museum, Departmento de its base, including the anal lobe, and mounting on
Biologia, Pontificia Universidad Catolica del Ecuador, card with water-soluble glue. These were pinned
Quito, Ecuador); QPIM (Queensland Department of below the specimen, above the labels.
Primary Industries and Fisheries, Mareeba, Queen- Genitalia preparations: Male genitalia were
sland, Australia); RMNH (Nationaal Natuurhis- extracted from at least one male from each disparate
torische Museum [formerly Rijksmuseum van locality, amounting to at least ten specimens per
Natuurlijke Historie], Leiden, Netherlands); SANC species. The aedeagus and (for both males and
(South African National Collection of Insects, Plant females) the genital segment were cleared in KOH
Protection Research Institute, Pretoria, South Africa); and studied in glycerine. There has been much dis-
SEMC/KSEM (Snow Entomological Museum, Univer- cussion surrounding the most effective method of
sity of Kansas, Lawrence, KS, USA); ShibataY (Yasu- internal sac eversion (see Skelley, 1993 for review)
toshi Shibata private collection, Tokyo, Japan); TMSA and this is an unpredictable process. To evert the
(Transvaal Museum, Pretoria, South Africa); UQIC internal sac, aedeagi were first cleared, and then
(University of Queensland, St. Lucia, Australia); forceps were used in a push–pull combination that
USNM (National Museum of Natural History, Wash- simultaneously applied light pressure to the basal
ington DC, USA); UZIU* (Zoological Museum [= Evo- bulb while pulling on the copulatory piece. In some
lutionmuseet, Zoologi], Uppsala University, Uppsala, cases, a minuten pin was used to push internal struc-
Sweden); WADA (Western Australia Department of tures through the median orifice. In rare cases, basal
Agriculture, Perth, Australia); WAMP (Western Aus- bulb pressure produced abrupt and complete infla-
tralian Museum, Perth, Australia); ZMHB (Museum tion, which might best indicate what the fully inflated
für Naturkunde der Humboldt-Universität, Berlin, internal sac would look like in life (e.g. see Fig. 18H).
Germany); ZMUC (University of Copenhagen, Zoo- Female internal genitalia were extracted by separat-
logical Museum, Copenhagen, Denmark); ZMUM* ing the terminal abdominal tergites and sternites
(Zoological Museum, Moscow State University, (VI–VIII) and removing the genital segment while all
Moscow, Russia). structures were visible. Once cleared, the gonocoxites,
Specimens were labelled with a unique Field surrounding membrane, and common oviduct were
Museum Division of Insects 10-digit identifier removed to expose vaginal and other structures. All
number (quoted as ‘FMNH-INS 0000 0XX XXX’) when genitalia were stored in white-stopped polyethylene
no other equivalent identifiers were already present. genitalia vials pinned with the specimen.
The taxonomic revision incorporates 9550 specimens Measurements. When possible, at least ten male
of (mostly) the North American Creophilus maxillosus and ten female specimens were measured for each
728 D. J. CLARKE
species using a calibrated eyepiece ocular micrometer. ‘. . . the first label, just beneath the specimen, is the
These specimens covered the size range of each label linked to the specimen. The other ones are
species, and included the largest and smallest speci- localities from where Fauvel saw specimens’ (D. Drug-
mens. Measurements are reported in millimetres. mand, IRSNB, in litt., 9.ix.2005). The C. erythroceph-
Comparisons of morphometric characters, such as alus and C. oculatus type material in the Banks
head shape and length, in different species are made Collection (BMNH) is associated with ‘Type’ labels
with reference to specimens of similar sizes. Abbre- handwritten by C. O. Waterhouse and placed to the
viations for measurements are as follows: ESL, right side of any specimens and above the cabinet
elytral sutural length, taken from scutellar apex to name label. These labels appear to indicate the pres-
sutural apex; EW, elytral width; OL, maximum eye ence of syntypes in the modern sense and do not
length (ocular length); HL, head length measured indicate that one or other of the specimens has been
from frontal margin to nuchal groove; HL1, distance designated as a lectotype (R. Booth, BMNH, in litt.,
from frontal margin of head to imaginary line drawn 12.vi.2007). Further details on this material are given
at posterior margin of eyes; HL2, distance from that in the species treatments. The following conventions
line to the nuchal groove (see Fig. 2B); HW, maximum are used for presenting type specimen label data:
head width; IOW, maximum distance between eyes quotes (‘’) enclose label data quoted verbatim, a slash
(interocular width); ML, left mandible length mea- (/) separates different labels, a pipe (|) separates
sured from apex to mandibular articulation; ML1, different lines of the same label, square brackets []
distance from left mandible apex to anterior-most enclose my comments, and a semicolon (;) separates
tooth; ML2, distance from anterior-most tooth of left specimens or specimen lots with different label data.
mandible to mandibular articulation (see Fig. 2D); If the type locality was given in the original descrip-
PL, pronotal length; PW, maximum pronotal width. tion it is quoted verbatim and enclosed in quotes,
Wing length was taken as the greatest length, from otherwise it is inferred from the scope of the publi-
the base of the subcosta anterior bulge (Kukalová- cation and given without quotes. In accordance with
Peck & Lawrence, 1993) to the apex. To account for the ICZN (Article 76), the locality of a newly desig-
the telescoping of the abdomen in rove beetles, I use nated lectotype specimen is here designated as the
‘forebody’ length as a standardized proxy for body type locality of the species.
length (= HL + PL + ESL), and report this as the To facilitate comparison of similar species in the
range for each species. Standard measurement ratios Taxonomy section, species are grouped into their
for each species are reported in supporting Tables S4 respective species-groups, listed according to their
and S5. overall similarity and beginning with the ‘type-
species’ of each group. To facilitate easier cross-
referencing between the identification key and the
TYPE MATERIAL, TAXONOMIC CONVENTIONS, species treatments, the species are numbered 1–12 in
AND DISTRIBUTION DATA both places. Herman (2001b) was used as the primary
Primary type material for most names was borrowed literature source for this study and the reader is
for study, although many type specimens of C. max- referred there for a comprehensive list of generic
illosus synonyms could not be located. The status of citations and other literature. For C. maxillosus and
all type material was checked against the original its synonyms, only the following literature is given:
publications. The existence of a holotype for any (1) any known citations not in Herman (2001b), and
nominal species-group taxon was confirmed only if it (2) first synonymization and subsequent status
could be unambiguously established that the original changes.
description was based on a single specimen, or if the Specimens with distinct locality information but
author unambiguously indicated that a single speci- without geographical coordinates on labels were
men from a larger type series was the ‘type’. In cases georeferenced using various gazetteers, topographic
where a type series comprised syntypes, a lectotype maps, and original publications reporting species
was designated to stabilize the application of names. occurrences (see supporting information, Appendix
This was especially important, not only because of S1). For localities including a distance offset (e.g.
the many junior synonyms, but because historically 16 km north-west of Auckland) I used the MaNIS/
intraspecific size variability and corresponding mor- HerpNet/ORNIS georeferencing calculator available
phological (especially morphometric) differences were at http://manisnet.org/gc.html. Nearly all localities for
used to erect several new Creophilus species. Type C. maxillosus villosus were georeferenced by A. F.
material for unavailable names is listed but given in Newton using gazetteers or topographic maps;
quotes (e.g. ‘Syntypes’). Type specimens of some A. neither of us attempted to gather all records of C.
Fauvel species, and sometimes specimens studied by maxillosus, and thus do not provide a complete dis-
him, have multiple locality labels. In these cases tribution map for that species. Distribution maps
were generated in ArcMap 9.1 (ESRI) using shape autapomorphy of Hp. fossor, applies to both male and
files freely available mostly through the DIVA- female genitalia and so is not grouped with either of
GIS application of Biogeomancer at http://www. these systems. Species polymorphic for a character
biogeomancer.org/software.html. Shape files for New were assigned all relevant states. Scoring of the data
Zealand and the Chatham Islands were sourced from matrix is based entirely on observation of specimens.
Land Information New Zealand, via Koordinates
(http://koordinates.com/). Two-letter area codes for
New Zealand localities follow Crosby, Dugdale & Watt TAXA
(1998); for all other countries, with few exceptions,
The ingroup comprised all known species here con-
second-order administrative divisions are those listed
sidered part of the Creophilus complex, including the
in the US Federal Information Processing Standard
New Zealand endemic species Hadrotes wakefieldi,
Publication 10-4 ‘Countries, Dependencies, Areas of
considered by some authors to be misplaced (Kli-
Special Sovereignty, and Their Principal Administra-
maszewski, Newton & Thayer, 1996). In agreement
tive Divisions’ (FIPS PUB 10-4).
with recent treatments (e.g. Smetana & Davies, 2000;
Smetana, 2004), C. maxillosus is treated as having
PRIMARY HOMOLOGY ASSESSMENT, CHARACTER two distinct subspecies, and to test the monophyly of
SELECTION, AND CODING this species, I included them as separate terminals.
Selection of appropriate outgroups was problematic
Hypotheses of primary homology were represented
because although Creophilus is placed in Sta-
here as data matrix columns (following Hawkins,
phylinina, there is no existing broad phylogenetic
Hughes & Scotland, 1997), and character state iden-
hypothesis for this subtribe. Likewise, the sister-
tity (sensu Brower & Schawaroch, 1996) was inferred
group of the Creophilus complex is unknown.
by applying standard criteria of homology (Remane,
Outgroups were therefore selected as generalized
1952; Patterson, 1982). Phylogenetically informative
exemplars from a group of genera traditionally affili-
characters were sampled from the external body and
ated with Staphylinus Linnaeus, 1758 (see Smetana
from male genitalia (including internal sac) and
& Davies, 2000): Platydracus maculosus (Graven-
female genitalia. No phylogenetically informative
horst, 1802), Dinothenarus badipes (LeConte, 1863),
internal skeletal characters were observed. Larvae of
Ontholestes cingulatus (Gravenhorst, 1802), and Sta-
most species in the Creophilus complex are unknown,
phylinus caesareus Cederhjelm, 1798 (representing
and so no larval characters were included. Characters
Staphylinus sensu Smetana & Davies, 2000). Xantho-
were selected based on observable interspecific vari-
pygus xanthopygus (Nordmann, 1837) (subtribe
ability that could be parsed into two or more discrete
Xanthopygina) was also included, which seemed
or at least non-overlapping states. I surveyed
appropriate as Creophilus historically has been
related genera (most in subtribe Staphylinina), rel-
placed by some authors in Xanthopygina. Monophyly
evant monographs, and published phylogenies of
of Creophilus + Platydracus Thomson, 1858 + Dinoth-
higher staphylinine taxa in order to facilitate polarity
enarus Thomson, 1858 was supported in recent
assessment and detection of plesiomorphic ingroup
analyses of the tribe Staphylinini (Solodovnikov &
character states. This helped a priori to avoid
Schomann, 2009), but Xanthopygus Kraatz, 1857 was
interpretation of symplesiomorphy as synapomorphy
inferred to be distantly related to this group.
(‘symplesiomorphy trap’; Wägele, 2005), and based on
this survey, characters inferred to be plesiomorphic
for the ingroup were excluded from analyses. Char-
acters were constructed using reductive ‘coding’ PHYLOGENETIC ANALYSES
(sensu Wilkinson, 1995; see his paper for details), but PAUP* 4.0b10 (Swofford, 2002) was used for all phy-
it should be noted that this is not synonymous with logenetic analyses. All characters were unordered and
binary presence/absence coding (contra Strong & Lip- equally weighted and polarity was established using
scomb, 1999; Forey & Kitching, 2000). Morphological the outgroup criterion with the root set as Staphyli-
data were compiled in Mesquite (Maddison & Maddi- nus caesareus (listed first in the matrix). Parsimony-
son, 2009) and the complete data matrix (Appendix 1) uninformative characters were excluded from all
is divided into four character systems (process parti- analyses and calculations. It was possible to use the
tions of some authors). External structural characters branch-and-bound (implicit enumeration) algorithm
are frequently abbreviated as simply ‘structure’ (or for most analyses, and thus the discovery of minimum
‘S’; characters 1–48), male genitalia as ‘male’ (or ‘M’; treelength was guaranteed for the combined, struc-
characters 49–77), female genitalia as ‘female’ (‘F’; tural, and male genitalia matrices (bandb option
characters 79–91), and macrosetal characters as ‘chain PAUP* using default settings). Heuristic tree
etotaxy’ (or ‘C’; characters 92–130). Character 78, an searches were used for female genitalia and chaetot-
730 D. J. CLARKE
axy (1000 random addition sequence replicates, TBR Gatesy, 2002): (1) number of parsimony-informative
branch swapping to completion, maxtrees set to characters (PICs), (2) number of most-parsimonious
auto-increase by 100). Each character system was trees (MPTs), (3) tree length (TL), (4) consistency
analysed separately, and then combined in a simul- index (CI), (5) retention index (RI), (6) resolution
taneous analysis of all data (121 parsimony- (total number of nodes), and (7) number of nodes in
informative characters). Results of separate analyses each separate analysis congruent with the simulta-
were also summarized using a consensus tree (taxo- neous analysis topology. To compare statistics among
nomic congruence: Mickevich, 1978). separate and simultaneous analysis trees, I also cal-
culated the average TL, CI, and RI of each character
system on the MPTs for the simultaneous analysis
SUPPORT MEASURES
using the ‘Values for current tree’ and ‘Step through
Bootstrap proportions (Felsenstein, 1985), Bremer trees’ facilities in Mesquite.
support (Bremer, 1988), and the number of Partitioned support measures the contribution of
unambiguously optimized and uniquely optimized different character systems to the overall support for
synapomorphies were calculated to quantify nodal a hypothesis (i.e. for both an individual clade and
support/stability. Bootstrap proportions (BP) were the entire tree), and reveals the distribution of
calculated using PAUP* with heuristic search set- support across the tree. To quantify the relative con-
tings as described above. Bremer support (BS) was tribution of each character system to nodal support
calculated using TreeRot (Sorenson & Franzosa, measures (BS, SYN, and UNIQ), I calculated parti-
2007), and the output was visualized in FigTree tioned Bremer support (PBS; Baker & DeSalle, 1997)
(Rambaut, 2009). Character state changes were using TreeRot, partitioned synapomorphy (PSYN;
examined and mapped on the consensus tree for Gatesy et al., 1999), and partitioned uniquely opti-
each analysis using Winclada (Nixon, 2002) and mized synapomorphy (PUNIQ; this paper). PBS
Mesquite (Maddison & Maddison, 2009). Most values are negative when the character system
ambiguously optimized states were resolved to pre- favours the shortest trees without the given node
serve the strongest hypothesis of primary homology (Baker & DeSalle, 1997; Gatesy et al., 1999). PSYN
(e.g. see De Pinna, 1991); although in most cases and PUNIQ were compiled for each node and char-
this involved accepting the ACCTRAN (or fast) opti- acter system from the Winclada output as described
mization over the DELTRAN (or slow) optimization, above, and can be only positive or zero. For each of
this need not be so (Agnarsson & Miller, 2008). In PBS, PSYN, and PUNIQ, the sum of partitioned
all cases, secondary loss was therefore favoured over values at a node equals the nodal support value for
parallel gains of a structure. Character state that node (e.g. SPBS at a node = BS for that node:
changes with multiple possible state assignments at Baker & DeSalle, 1997). I report the totals of PBS,
a node were excluded from the preferred tree. Also, PSYN, and PUNIQ for the simultaneous analysis
only optimizations stable across all of the most- tree to record the contribution of each character
parsimonious trees are plotted and discussed. The system to the overall support for the simultaneous
Winclada output was used to tally unambiguous analysis tree (e.g. SPBS for all nodes = contribution
synapomorphies (SYN) and uniquely optimized of each character system to overall BS: Baker et al.,
synapomorphies (UNIQ) supporting nodes of the 2001). The sum of these summed values is equal to
consensus trees for each analysis. Only those the total support for the tree, measured as total BS
present in all most-parsimonious trees in an analy- (Källersjö et al., 1992), SYN, or UNIQ. To account for
sis were recorded, and this was determined using differences in dataset size (numbers of characters
the ‘Trace character over trees’ facility in Mesquite. and states), each measure of total support was also
This ensured that character support for the pre- standardized by a relevant quantity [SPBS/minimum
ferred hypothesis, as well as calculations involving number of steps required for a character system
SYN and UNIQ values, were based only on charac- (Baker et al., 2001); SSYN/total number of character
ter state changes supported by all possible resolu- states for a character system; SPUNIQ/SPSYN, total
tions of soft polytomies, and thus results are number of unambiguous synapomorphies for a char-
independent of any particular preferred tree. acter system].
Hidden support, or secondary signal emerging in
a simultaneous analysis, is a critical aspect of the
ASSESSING THE PHYLOGENETIC UTILITY OF relative phylogenetic utility of different character
CHARACTER SYSTEMS systems (Chippindale & Wiens, 1994; Nixon & Car-
To compare different character systems, I used penter, 1996; Gatesy et al., 1999). To quantify
several widely used metrics of ‘dataset quality’ (e.g. hidden support, I calculated hidden Bremer support
de Queiroz & Wimberger, 1993; Baker et al., 2001; (HBS; Gatesy et al., 1999), hidden synapomorphy
(HSYN; Gatesy et al., 1999), and hidden uniquely anteriorly for some distance beyond the transverse
optimized synapomorphy (HUNIQ; this paper). part of the postgenal ridge (Fig. 2A, io). Other
Hidden support values can be positive, zero, or ridges are variable among or within Creophilus
negative, whether measured as HBS, HSYN, or species, particularly the medial fate of the postgenal
HUNIQ. To quantify the relative contribution of ridge [obsolete mesially in C. albertisi (Fauvel,
each character system to the hidden support (mea- 1879), Fig. 31B, pgr] and the dorsal completeness of
sured as BS, SYN, and UNIQ) at each node, I cal- the nuchal ridge. The latter is usually incomplete
culated hidden partitioned Bremer support (HPBS), dorsally, such that the posterior part of the vertex is
hidden partitioned unambiguous synapomorphy continuous with the disc of the neck medially
(HPSYN), and hidden partitioned uniquely opti- (Fig. 2B, nr), but is also intraspecifically variable
mized synapomorphy (HPUNIQ). For a given node (hence excluded from analyses). It is always com-
and character system in the simultaneous analysis, plete in Liusus. The postmandibular ridge is absent,
hidden partitioned support is the difference between but one new character of the erythrocephalus-group,
the partitioned support value for a character system the ‘ventrolateral carina’ (ch6-1; Fig. 2A, vl) of large
– PBS, PSYN, or PUNIQ – and the support value males, may be homologous to the basal part of the
for that node in the separate analysis of that char- postmandibular ridge. [Note: this ridge is somewhat
acter system – BS-sep, SYN-sep, or UNIQ-sep. (e.g. obliterated by numerous confluent punctures in C.
HPBS for node A = PBS for node A – BS-sep for lanio (Erichson, 1839) and is present in all individu-
node A). The hidden support at a node then is the als of C. albertisi, in which it is distinctly modified
sum of partitioned hidden support values, and total into an anterolaterally projecting tooth; Fig. 31A, B,
hidden support emergent in the simultaneous analy- vl.]
sis is the sum of hidden support values at each The configuration of the superior and inferior mar-
node. For more complete details and explanation ginal lines of the pronotal hypomeron (ch22; Fig. 2E,
of support measures used here, see Gatesy et al. il, sl) was recently clarified for several staphylinine
(1999). genera (Solodovnikov & Schomann, 2009) in that the
conditions of ‘separate’ vs. ‘joined’ can conceal several
qualitatively different states. With the exception of
REVIEW OF MORPHOLOGICAL Thinopinus pictus LeConte, 1852, which is polymor-
CHARACTER SYSTEMS phic for this character, the Creophilus complex
References to character numbers (e.g. ‘ch65-1’) refer (including Ht. wakefieldi) and X. xanthopygus each
to those in the list of phylogenetic characters. have the inferior marginal line separate from the
superior line and continuous with the anterior line
(ch22-1). Hadrotes wakefieldi and X. xanthopygus,
EXTERNAL MORPHOLOGICAL CHARACTERS however, differ from other Creophilus complex
(STRUCTURE, S; CHARACTERS 1–48) species in having the superior line widely separate
These characters include all external cuticular (and anteriorly from the inferior line. A new character for
other) features of the body and appendages, except Staphylinini reported here is the presence of distinct
macrosetae. General terminology follows Blackwelder ridges on the mesepimeron (ch27). In all outgroups
(1936) and Smetana & Davies (2000), with some there are two distinct ridges (Fig. 9I, arrows),
terms updated to those used by Thayer (2005). Wing whereas all Creophilus complex species have only a
terminology follows Kukalová-Peck & Lawrence single ridge at the edge of the ventral sclerite
(1993). I use ‘postgena’ to refer to the ventral side of margin (Fig. 9K, arrow). A carina not previously
the head. recorded for this group is the secondary basal trans-
Cuticular ridges. The homology of several head verse carina on abdominal segments III–V (ch45-1)
ridges is often questionable within Staphylinini (e.g. in the erythrocephalus group (Fig. 2L, sbr; VI also in
see discussions in Solodovnikov, 2005; Solodovnikov Ht. wakefieldi). Similar ridges are found throughout
& Newton, 2005), but Smetana & Davies (2000) is Staphylininae.
followed for names of various ridges and carinae. In Mandibles. Mandibular teeth have been shown to
Creophilus, the configuration of the ventral basal, be taxonomically informative within Staphylinini
postgenal, infraorbital, nuchal, and dorsal basal (e.g. Smetana & Davies, 2000, and references
ridges is more or less exactly as figured by Smetana therein). The dorsal teeth (labelled T1 and T2) and
& Davies (2000: fig. 2), with some exceptions. The ventral tooth (T3) of the right mandible are consid-
infraorbital ridge is present (contra Solodovnikov & ered homologous among taxa (Fig. 2D). T2 is also
Schomann, 2009) basally as a ventrally curving the medial termination of the mandibular blade
short ridge lying approximate to the lateral portion (ch14-1), which is absent in Hp. fossor and T. pictus
of the postgenal ridge, and in most species extends (ch14-0). Given that the two teeth in these species
732 D. J. CLARKE
are in the same plane, it is hypothesized that they and ventral to T2 in all seven species (topographic
are homologous to T1 and T2 in other species. Three and fine structural criteria), it is hypothesized that
erythrocephalus-group species (C. erythrocephalus, these two teeth are homologous to T2 and T3. Ven-
C. imitator Cameron, 1952, and C. albertisi) have trally, the left mandible in Creophilus has a distinct
only two distinct teeth on the right mandible. Based basolateral ridge (ch13-1; Fig. 2C, vbr) that is
on the observation that the largest tooth is proximal always best developed in large males, although fre-
Figure 2. Creophilus, details of morphology (setae omitted except in H–K). C. erythrocephalus (A-B, L, large male); C.
variegatus (C, D, large male); C. maxillosus maxillosus (E–G); C. lanio (H–J); C. incanus (K). A, head, ventral (mouthparts
removed); B, head, dorsal (antennae removed); C, left mandible, lateral and ventral (prostheca omitted); D, mandibles,
dorsal (prostheca omitted); E, prothorax, ventral (small male); F, pterothorax, ventral; G, pronotum, dorsal (large male);
H, left maxilla, ventral; I, labium, ventral; J, abdominal tergite II, dorsal; K, right protibia, anterior (smaller setae/bristles
omitted); L, abdominal segments III–V, dorsal. Abbreviations: af, anterolateral flange of abdominal tergite II; atf, antennal
fossa; bgi, basal gular impression; bpd, basolateral pronotal denticle; btf, basal transverse fold; dbl, dorsal basolateral
ridge; dbr, dorsal basal ridge; f, muricate fossa of abdominal tergite II; HL, head length; il, inferior pronotal line; io,
infraorbital ridge; mac, posterior marginal carina of mesocoxal acetabulum; mb, mandibular blade; mip, mesoventral
intercoxal projection; ML, mandible length; mt, medial tumescence of abdominal tergite II; nr, nuchal ridge; pf,
posteromedian furcasternal fovea; pg, postgenal ridge; plf, posterolateral fossa; pmg, premetacoxal groove; pos, postoc-
cipital suture; ppp, pronotal postcoxal process; ps (fused) parasclerites of abdominal segment II; ptm, mesothoracic
peritreme; sbr, secondary basal ridge; sl, superior pronotal line; spi, spiracle; sr, sternacoxal ridge; T1, T2, T3, mandibular
teeth; td, transverse depression of abdominal tergite II; vb, ventral basal ridge; vbr, ventral basolateral ridge of left
mandible; vl, ventrolateral carina. Scale bars = 1 mm.
䉳
quently variously effaced in some species and Fig. 3M; except in C. variegatus), supplied with
smaller specimens. When closed, the left mandible several moderately long setae, and never extends
is dorsal to the right mandible and the groove beyond the median lobe apex.
created by the ventral basolateral ridge receives the Internal sac. In all terminal taxa the internal sac
inner edge of the right mandible. everts symmetrically, without twisting (as noted for
many related species by Coiffait, 1956), and all
structures are bilaterally symmetrical about the lon-
MALE GENITALIA (MALE, M; CHARACTERS 49–77) gitudinal median lobe apex. The most conspicuous of
Male genitalia include abdominal segments IX and the sclerotized internal sac structures is the elon-
X (see Blackwelder, 1936) and the aedeagus. The gate copulatory piece (Fig. 3, cp; cane-like in most,
aedeagus of Creophilus maxillosus (including some chs70-1 and 71-1; straight in T. pictus and Hp.
internal sac details) was described by Blackwelder fossor; ch71-0, Fig. 3E). When everted, this structure
(1936) and Coiffait (1956), and in detail by Dajoz & lies in a plane either subparallel to the median lobe
Caussanel (1969). axis (Fig. 3D, E) or subperpendicular to it (Fig. 3H).
Median lobe and paramere. The question of which The latter suggests that in those species the median
side of the aedeagus is morphologically dorsal has lobe apex does not enter the female during copula-
been discussed by several authors (e.g. see Gusarov, tion. The copulatory piece is either surrounded by a
2002). I follow Thayer (2005), and refer to the median large spiculose membranous sheath (Figs 3, 4, sh),
foramen (parameral) side as dorsal, an interpretation or not (e.g. C. variegatus; Fig. 4A). The appendix of
supported by pupal development in Philonthus the copulatory piece (ch73; hereafter ‘appendix’;
(Tikhomirova, 1978). (Note: in the Creophilus complex Figs 3, 4, app) varies interspecifically in its connec-
the aedeagus in repose lies with the parameral side tivity with the copulatory piece. Its structure is
facing left.) In the erythrocephalus-group and Liusus, usually species-specific, and may be important for
the apical part of the median lobe is continuously mechanical recognition during copulation. Attached
sclerotized ventrally (ch52-0; Fig. 3A), whereas in all basally to the copulatory piece is the internal ‘basal
other species there is a non-sclerotized strip connect- inner tube’ (ch62-1; Figs 3, 4G, bit), an apparently
ing the median orifice with the basal bulb (ch52-1; muscular structure perhaps functioning as a sperm-
Fig. 3C, arrow). The interspecific variability in the pump. The ‘basal cap’ (ch64-1; Figs 3, 4G, bc) is a
shape of the median lobe apex indicates that the apex round sclerite at the base of the inner tube through
may be related to internal sac inflation during copu- which the ejaculatory duct passes. It may serve
lation, or to female genital stimulation. At least in C. either as a muscle attachment point for internal sac
maxillosus, the apex of the median lobe actually retraction and/or basal inner tube contraction. The
enters the female during copulation (pers. observ.), ejaculatory duct outside the median lobe comprises
which may not be the case for erythrocephalus-group a distinctly sclerotized section near the median
species (see below). Except for Ht. wakefieldi and C. foramen (absent in O. cingulatus, P. maculosus, and
albertisi, in which it is fused basally to the median Ht. wakefieldi). I could not establish that this was a
lobe, the paramere in all species is articulated at its homologous structure in different genera, and it
base to the basal bulb and is dorsoventrally movable requires further study. The ‘ventral process’ (ch68;
(ch59-1). The parameral apex is notched (ch61-1; Fig. 3, vp) is reduced in the erythrocephalus-group
734 D. J. CLARKE
Figure 3. Male genitalia, details of morphology. Creophilus erythrocephalus (A, B, G, H, O); C. lanio (Q); C. maxillosus
maxillosus (C, D, I, P); C. oculatus (J); Hadropinus fossor (E, F); Staphylinus caesareus (K, L); Hadrotes wakefieldi (M);
Dinothenarus badipes (N). A, aedeagus with details of inverted internal sac, ventral; B, same, right lateral (parameral
setae omitted); C, median lobe with fully everted internal sac, ventral – arrow indicates membranous region; D, apical
part of median lobe with fully everted internal sac, right lateral; E, aedeagus with fully everted internal sac, right lateral
(parameral setae omitted), note: bit has become separated from cp in this specimen; F, copulatory piece and appendix of
internal sac, dorsal; G, ventral sclerite of internal sac, ventral; H, apex of median lobe with fully everted internal sac,
right lateral; I, ventral sclerite of internal sac, ventral (lower) and apical (upper); J, apical part of median lobe with
partially everted internal sac, ventral; K, apical part of median lobe showing part of everted internal sac, right lateral;
L–N, paramere, dorsal; O–P, median lobe apex, ventral; Q, male genital segment (setae omitted). Abbreviations: app,
appendix; as, apicolateral sclerite of median lobe; bc, basal cap; bit, basal inner tube; bms, basomedian sclerite of internal
sac; ce, carinate edge of median orifice; cp, copulatory piece; dr, dorsal receptacle for copulatory piece/appendix; is, internal
sac; lit, lateral inflexed tooth of median lobe; ml, membranous lobe; mlb, median lobe of aedeagus; stm, subtriangular
membranous region; vp, ventral process; vr, ventromedian spiculose strip; vs, ventral sclerite; sh, spinose sheath
surrounding copulatory piece; sr, one of pair of basoventral spiculose regions. Scale bars: G and I = 0.1 mm; rest = 1 mm.
䉳
Figure 4. Male copulatory pieces of selected species in the Creophilus-complex, dorsal view. In each figure the large
paired dark structures are the sclerotized arms of the copulatory piece. A, C. variegatus; B, C. maxillosus maxillosus; C,
C. m. villosus; D, C. galapagensis; E, C. flavipennis; F, C. incanus; G, C. oculatus; H, C. erythrocephalus; I, C. imitator;
J, C. lanio; K, C. albertisi; L, Hadrotes crassus; M, Thinopinus pictus; N, Hadropinus fossor; O, Liusus hilleri.
Abbreviations: iedg, apical inner edge of arms of copulatory piece; por, parallel oblique ridges of copulatory piece (see also
Fig. 10); xbar, ‘cross-bar’ of appendix. Additional abbreviations as in Figure 3. Scale bar = 1 mm.
736 D. J. CLARKE
Figure 5. Female genitalia, details of morphology (all but E, ventral view). A, Creophilus m. villosus, internal genitalia;
B, C. erythrocephalus, internal genitalia; C, C. huttoni, internal genitalia; D, C. oculatus, gonocoxites and styli (setae
omitted from left side); E, Thinopinus pictus, abdominal segments VIII–X, dorsal (vestiture omitted); F, T. pictus, internal
genitalia; G, Staphylinus caesareus, internal genitalia; H, Platydracus maculosus, internal genitalia; I, Hadrotes crassus,
right gonocoxites and stylus (vestiture omitted) – arrow indicates membranous articulation; J, T. pictus, right gonocoxite.
Abbreviations: bcx, bursa copulatrix; fl, flagellum (note that the complete structure is not shown); gc, gonocoxite; gcm,
gonocoxal macroseta; gc1, gonocoxite I, gc2, gonocoxite II; mg, median groove; pls, paired lateral sclerites of vaginal plate;
sc, sclerite of bursa copulatrix; sd, spermathecal duct; sp, spermatheca; ss, sclerotized strip of vaginal plate; sty, gonostyle;
vf, vaginal fold; vp, vaginal plate. Scale bars = 1 mm.
(ch68-2; Fig. 3H) such that the ‘ventromedian spicu- FEMALE GENITALIA (FEMALE, F; CHARACTERS 79–91)
lose strip’ (ch67-1, Fig. 3, vr) lies nearly flat on the
surface of the everted internal sac (Fig. 3H), subad- Female genitalia include abdominal segments IX and
jacent to the paired ‘basoventral spiculose regions’ X (including the gonocoxites: Fig. 5D, E, I, J, gc),
(ch66-1; Fig. 3, sr). The shape of the ventral sclerite internal vaginal structures, bursa copulatrix, and
(ch69-1; Fig. 3, vs) is species-specific. The dorsal spermatheca (Tuxen, 1970). Tanner (1927) and Black-
receptacle for the apex of the copulatory piece welder (1936) described only external structures of C.
(ch76-1; Fig. 3, dr; Blackwelder, 1936: fig. 9G, z) is maxillosus, whereas Dajoz & Caussanel (1969) and
apically grooved for receiving the copulatory piece De Marzo (2010) described internal structures. I
apex/appendix when the internal sac is inverted. mainly follow the terminology of Dajoz & Caussanel
(1969), but note that their ‘spermathèque’ [spermath- ogy across these studies.) Macrosetae identified in
eca] is labelled ‘spermathecal gland’ by De Marzo this study are shown in Figure 6. Topography is espe-
(2010). Illustrations in Dajoz & Caussanel (1969: cially important for chaetotaxy, and permits recogni-
fig. 10) provide the anatomical context for internal tion of ‘macrosetal fields’ (e.g. head: temporal and
characters. occipital macrosetae; elytra: ‘discal series’ and ‘poste-
The ‘vaginal plate’ on the dorsal surface of the rior elytral patch’; legs: setal patches such as the
vaginal cavity is as diagnostically valuable as the ‘procoxal patch’; abdomen: additional tergal macrose-
aedeagus and contributes phylogenetically informa- tae in Ht. crassus).
tive characters (ch85-0, Fig. 9N, middle arrow, O; In Creophilus there is a generally clear pattern of
ch85-1, Fig. 9M; chs86-1 and ch86-2, Fig. 5B, C, pls). serially homologous abdominal tergal macrosetae
The bursa copulatrix (ch88-1; Fig. 5, bcx) is delimited (Fig. 6, il, mm, ol). I use a six-digit number (‘tergal
ventrally by a large (and in Staphylinus very large) chaetotaxic formula’) to describe the macrosetae of
‘sclerotized plate’ embedded in the vaginal cavity abdominal tergites III–VIII. Polymorphic variation
wall/common oviduct (ch87-1; Fig. 5, sc, and Fig. 9N, is recorded as numbers in braces. For example, the
left arrow, O). Immediately posterior and adjacent to tergal chaetotaxic formula 6–6-6-4(6)-6-8(12) should
the vaginal plate apex is the ‘vaginal fold’, which is be interpreted as all six macrosetae present on terg-
variably sclerotized in the erythrocephalus-group ites III–V and VII, only four (occasionally six) on
(chs82-1, 2, and 3; e.g. Fig. 5B, C, vf), but membra- tergite VI, and eight (occasionally up to 12) on
nous or scaled in other species (ch82-0, Figs 5A, G, tergite VIII. There are often ‘extra’ macrosetae (par-
9N, right arrow, Fig. 16I, vf). All structures anterior ticularly in C. flavipennis and Ht. crassus) in a mac-
to the bursa copulatrix opening are within the body rosetal field (and frequently on the abdomen), but
cavity. Anteriorly, the bursa copulatrix opens into an these are never consistently present and rarely
enlarged pouch-like cavity and is extended into a bilaterally symmetrical (see also Smetana, 1971 and
gradually attenuating tubular structure (Fig. 5, bcx). Smetana, 1995 for Nearctic Quedius and Philonthus,
During copulation, the male copulatory piece/ respectively). For phylogenetic analyses, abdominal
appendix complex probably enters this structure. tergal macrosetae could have been coded as follows:
Continuing anteriorly is an extremely long ‘flagel- (1) inner lateral, outer lateral, and medial macrose-
lum’ (ch90-1; Fig. 5, fl) that becomes bunched into a tae each considered as a separate character for each
ball-of-wool-like structure. If unravelled, the flagel- tergite (18 characters); (2) each of these considered
lum would be many times the length of the insect as serially homologous on each tergite and coded as
(see De Marzo, 2010: fig. 12D, RIC). The spermath- three multistate characters; or (3) all macrosetae
eca in most species is narrow and thread-like coded as a single complex multistate character. The
without distinct duct and reservoir (ch91-1; Fig. 5B, first method seemed most appropriate as most
C, sp). Hadrotes wakefieldi and P. maculosus have species have different combinations of macrosetae,
very different structures in that the spermatheca but also because polymorphisms would further com-
opens directly into the vaginal cavity, via an elon- plicate the coding of more composite characters. The
gate duct (ch89-1), and there is neither a bursa copu- question of character independence is complicated
latrix per se nor an associated flagellum (chs88-0, by interspecific patterns of variation, and so the 11
90-0; Fig. 5H). The data presented in this study informative characters used here represent a com-
suggest that further broad-scale, higher-level com- promise between assuming independence of all 18
parative studies are warranted. possible characters and the more conservative
combination of non-varying macrosetae on adjacent
tergites.
ADULT CHAETOTAXY (CHAETOTAXY, C;
CHARACTERS 92–130)
Broadly defined, chaetotaxy can include a variety of LIST OF CHARACTERS
mainly sensory structures that are traditionally An asterisk (*) indicates characters or character
important in larval systematics (see discussion in systems novel to this study, or characters apparently
Solodovnikov, 2007). In this study, chaetotaxy used phylogenetically for the first time.
includes only adult macrosetae – large setae in a
consistent location (Fig. 6), which are presumably
mechanosensory in function. Based on positional cri- EXTERNAL STRUCTURAL CHARACTERS
teria, all macrosetae that could be reliably homolo- (STRUCTURE, S)
gized are given names, mostly taken from Naomi Head
(1982), Hayashi (1993), and Campbell (1979). (Note: 1. General head integument colour (Newton, 1985):
use of these names does not necessarily imply homol- (0) black or brown, same as body integument
738 D. J. CLARKE
Figure 6. Scheme of taxonomically and phylogenetically

informative macrosetae (chaetotaxy) identified in this
study. A, head, prothorax, and pterothorax (dorsal); B,
same (ventral); C, abdomen, dorsal. Abbreviations: (Head)
– ad, additional temporal; al, anterolateral; am, antero-
marginal; ba, basiantennal; gn, genal; mn, mental; oc,
occipital; pl, posterolateral; sc, scapal; sm, submental; so,
supraorbital; tm, temporal; vs, ventroscapal; (Prothorax) –
an, anterolateral pronotal; ap, anterior pronotal group; bp,
basolateral pronotal; bs, basisternal; pc, apical procoxal
group; pp, procoxal patch; tr, protrochanteral; (Pterotho-
rax) – ae, anterior elytral; bh, basohumeral; ds, discal
series; hm, humeral; ma, metatrochanteral; mc, apical
mesocoxal; me, metacoxal patch; mo, apical metacoxal; mp,
mesocoxal patch; mr, mesoventral row; mt, mesotrochant-
eral; mv, metaventrital discal; pe, posterior elytral; pm,
posteromedian metaventrital; ps1, parascutellar 1; ps2,
parascutellar 2; pt, posterior elytral patch; (Abdomen) – il,
inner lateral; mm, median; ol, outer lateral; pa, paras-
cleral. III–VIII indicates numbering of true morphological
segments (I and II being reduced and hidden beneath the
elytra). The tergal chaetotaxy formula shown is
6–6–6–6–6–6.
䉳
(Fig. 1F–O); (1) bright orange-red or yellowish,

always at least orange-red or yellowish ven-
trally, and different from body integument
(Fig. 1A–E). Note that C. rekohuensis is only
darkened dorsally.
2. Secondary patterning of vertex/frons: (0) with
single medial black spot (Figs 1A, 2B, 7A, B,
D); (1) completely black between eyes
(Figs 1B–D, 7C; note the three New Zealand
species share the general state of having an
expanded darker pigmentation between the
eyes, Creophilus huttoni (Broun, 1880),
however, is often lighter and patchier overall,
as in Figure 1B, whereas C. rekohuensis has
the pigmentation additionally expanded behind
the eyes); (2) with variable black maculae
(Fig. 1M); (3) with triangular black spot
(Figs 1E, 7E, 31A); (4) absent. Thinopinus
pictus (scored state 2) is variable, with
pigment patterning varying over entire body
and within and between populations.
3. *Antennal fossae: (0) contiguous with frontocly-
peal margin, forming part of margin (Fig. 8A);
(1) separated from frontoclypeal margin by dis-
tinct sclerotized strip (Fig. 2B, af).
4. Punctation of vertex: (0) umbilicate punctures;
(1) small abrupt pits (Fig. 9C); (2) small
gradual pits (Fig. 9A); (3) absent such that
surface is smooth and shiny.
5. Head microsculpture: (0) absent; (1) anasto-
mosing pits and irregular grooves; (2) irregu-
Figure 7. Heads of some species of the Creophilus erythrocephalus species-group (antennal flagella omitted). A, C.
erythrocephalus; B, C. lanio; C, C. oculatus; D, C. imitator; E, C. albertisi. A–D, large males; E, small female. Scale
bars = 5 mm.
larly micropunctate (Fig. 9A, B); (3) subparallel 12. Longitudinal furrows laterad of mandibular
wavy lines. basolateral ridge: (0) absent; (1) present.
6. *Ventrolateral carina in large males: (0) absent 13. *Ventral basolateral ridge of left mandible:
(Fig. 8B); (1) present (Figs 2A, 30B, vl). Avail- (0) absent (Fig. 8B); (1) present (Fig. 2C,
able males of C. rekohuensis are smaller than vbr).
the minimum-sized males expressing state (1) 14. *Mandibular blade of right mandible: (0) absent
in other species, so this species is scored as (Fig. 1K, M); (1) present (Fig. 2D, mb).
unknown (?). 15. Mandibular tooth T1 of right mandible: (0)
7. Ventral basal ridge (Smetana & Davies, 2000): greatly reduced or absent (Figs 25B, 27A, 31C);
(0) contiguous with ventral portion of postoc- (1) present (e.g. Figs 2D, 13A).
cipital suture adjacent to basal gular impres- 16. Mandibular (ventral) tooth T3 of right mandible:
sion (Fig. 8B, bgi, vb); (1) separate from (0) absent; (1) present (Fig. 2D). Note that this
ventral portion of postoccipital suture adjacent tooth is located ventral to T2, regardless
to basal gular impression (Fig. 2A, bgi, vb). of whether the base of it is visible in dorsal view
8. Antennal structure: (0) antennomeres 7–10 with and thus slightly proximal to T2 (erythrocephalus
similar diameters; (1) clavate, antennomeres 7–10 group, Liusus) or whether it is partly to mostly
gradually increasing in diameter (e.g. Fig. 13F). obscured (maxillosus group), in which case it is
9. Antennal micropubescence distribution: (0) on directly beneath the mandibular blade.
antennomeres 4–11; (1) on antennomeres 5–11; 17. Largest tooth of right mandible in males: (0)
(2) on antennomeres 6–11; (3) on antennomeres all teeth approximately same size; (1) T1;
7–11. (2) T2; (3) T3; (4) T2 and T3 approximately same
10. Apical structure of antennomere 11: (0) produced size.
into anterior subconical point (e.g. Fig. 36B); (1) 18. Position of T3 on right mandible: (0) ventral to
produced into convex apical ridge (e.g. Fig. 16B); mandibular blade (Fig. 2D); (1) proximal to man-
(2) produced into emarginate apical ridge (e.g. dibular blade (but still in ventral plane; e.g.
Fig. 13F). Fig. 25B). See comment under character 16.
11. Structure of mandibular teeth: (0) small and 19. Vestiture of disc of head and pronotum: (0)
denticulate (Fig. 2C, D, T1–T3); (1) large and absent (Fig. 1); (1) present. The vestiture of
explanate (Fig. 1K, M). the head frequently co-varies with that of the
740 D. J. CLARKE
Figure 8. Details of morphology (vestiture omitted). Platydracus maculosus (A–C); Hadropinus fossor (D, E); Thinopinus
pictus (F, G). A, head, dorsal; B, head, ventral; C, abdominal segments II–V, dorsal; D, right protibia, posterior; E, same,
dorsal; F, abdominal segments II and III, dorsolateral; G, abdominal segment II, dorsal. Abbreviations: af, anterolateral
flange; bgi, basal gular impression; io, infraorbital ridge; mem, medial emargination; nr, nuchal ridge; pg, postgenal ridge;
po, postoccipital suture; ps, parasclerite; sbm, sinuate basal abdominal margin; spi, spiracle; vb, ventral basal ridge.
Abbreviations of macrosetae as in Figure 6. Scale bars = 1 mm.
pronotum in Staphylinini, so to avoid over- Davies, 2000); (1) not joining, inferior pronotal
weighting this variation it is combined into a marginal carina continuous with anterior prono-
single character. tal margin (Fig. 2E, il, sl, Fig. 9H, J).
23. Sternacostal ridge (Smetana & Davies, 2000):
Prothorax (0) projecting anteromedially (Fig. 9G, arrow); (1)
20. Basolateral pronotal impressions: (0) absent; (1) arcuate medially (Fig. 2E, sr, Fig. 9F, top arrow).
present, asperate, setose; (2) present, smooth, 24. Posteromedian fovea of furcasternum: (0) absent;
asetose. (1) present (Fig. 2E, pf).
21. *Basolateral pronotal denticle in large males: (0)
absent (Fig. 25A); (1) present (Fig. 2G, bpd). Pterothorax
Available males of C. rekohuensis are smaller 25. *Structure of mesothoracic peritremes: (0) small,
than the minimum-sized males expressing state with medial apex more or less rounded or trun-
(1) in other species, so this species is scored as cate; (1) very large, acutely produced medially
unknown (?). (Fig. 2E, ptm).
22. Superior and inferior marginal carinae of prono- 26. Shape of mesosternal intercoxal process: (0)
tal hypomeron: (0) joining behind anterior prono- truncate or broadly rounded (Fig. 2F, mip); (1)
tal angles, superior marginal carina continuous acuminate; (2) angulate (Fig. 33H, I); (3) nar-
with anterior pronotal margin (see Smetana & rowly rounded.
27. *Mesepimeron: (0) with two ridges (Fig. 9I, row of spines, and have two spines on the edge
arrows); (1) with single ridge at ventral edge of of the posterior face. Ontholestes cingulatus has
sclerite (Fig. 9K, arrow). 3–4 spines, not arranged into row(s), on the
28. Posterior marginal carinae of mesocoxal posterior face.
acetabula: (0) complete mesially; (1) obsolete 38. Apical protibial development: (0) normal,
mesially (Fig. 2F, mac). slightly expanded laterally, not produced into
29. *Intercoxal tumescence of metaventrite: (0) explanate projection (Fig. 2 K); (1) subfossorial,
absent; (1) present. This character comprises a posterior face convex; (2) fossorial, produced lat-
small mound at the apex of the metaventrital erally into explanate projection, posterior face
‘intercoxal process’. concave (Fig. 8D, E).
30. *Premetacoxal groove of metaventrite: (0) absent;
(1) present only laterally, obsolete mesially; (2) Abdomen
present, complete (Fig. 2F, pmg). The premeta- 39. *Medial tumescence of abdominal tergite II: (0)
coxal groove is present in all species of Creophi- absent; (1) present (Fig. 2J, mt)
lus as a distinctly wide and deep groove 40. *Anterolateral flanges of abdominal tergite II:
extending transversely in front of the meta- (0) strongly developed anteriorly, margin of
coxae, just before the posterior metaventrital tergite therefore distinctly emarginate (Fig. 8C,
margins. In state (1) the groove is distinctly af, mem); (1) poorly developed anteriorly,
effaced, and therefore broadly discontinuous, margin of tergite broadly rounded (Figs 2J, 8G,
mesially. af). This character describes the development
31. Central part of metaventrital disc: (0) setose and extent of internal extensions of the ante-
(Fig. 14); (1) glabrous. rior margin of tergite II, which in state (0)
32. Elytra when closed: (0) meeting at suture; (1) protrude somewhat creating a medial emargin-
overlapping. This character is an autapomorphy ation of the tergite.
of T. pictus. 41. Paired muricate fossae of tergite II (Dajoz &
33. *Distribution of elytral macrovestiture: (0) Caussanel, 1969: fig. 7A, Fs): (0) absent (Fig. 8G);
evenly covering elytra (Fig. 1M); (1) humeri at (1) present (Figs 2J, f, Fig. 9L, arrow). State (0) is
least partly glabrous (Fig. 1A–J); (2) marginal an autapomorphy of T. pictus).
only, disc glabrous (Fig. 1L, N, O); (3) parasu- 42. Spiracles of abdominal segment II: (0) in tergal
tural region of disc glabrous (Fig. 1K). notch surrounded by membrane (Fig. 2J, sp); (1)
34. Pattern of lighter coloured elytral macrovesti- embedded in tergite (Fig. 8G, spi). State (1) is an
ture: (0) absent, macrovestiture uniformly black autapomorphy of T. pictus.
or brown (Fig. 1A–E, H, M); (1) present (Figs 1F, 43. *Parasclerites of abdominal segment II: (0)
G, I, J, 13B, 16H, 23G). fused at least posteriorly, not separated by
35. Wing development: (0) fully developed, longer clear strip (Figs 2J, 8C, ps); (1) separated by
than elytra, functional as flying organ (e.g. distinct clear strip (Fig. 8F, G, ps). In all
Fig. 33N); (1) shortened, longer than elytra, not species scored (0), except C. galapagensis,
functional as flying organ (Fig. 33O, P); (2) there is a notch in the anterior margin of the
poorly developed, shorter than elytra, not func- sclerite. In T. pictus and Hp. fossor the two
tional as flying organ. sclerites are completely separated by a clear
strip.
Legs 44. Basal margins of abdominal tergites III–V: (0)
36. *Basal spine-like projection on profemur in large distinctly sinuate medially (Fig. 8C, sbm); (1)
males: (0) absent; (1) present (Fig. 9D, arrow). straight medially (Fig. 2L, btf ).
Available males of C. rekohuensis are smaller 45. Secondary basal transverse carina of abdomi-
than the minimum-sized males expressing state nal tergites III–V: (0) absent (Fig. 8C); (1)
(1) in other species, so this species is scored as present (Fig. 2L, sbr).
unknown (?). 46. Subtriangular posteromedial impunctate region
37. *Mesal pair of protibial spines: (0) absent; (1) of abdominal tergites III–VI: (0) absent, terg-
present (Fig. 9E, arrows). These spines are dis- ites evenly covered in vestiture; (1) present
tinctly separate from other (posterior) tibial (Fig. 1L, N).
spines. Outgroups have variously developed 47. *Apical fringe setae of abdominal tergites III–
protibial spines. Platydracus maculosus has an VI: (0) uniformly darkly coloured (e.g. Fig. 1A–
external row of 4–5 spines, with 1–2 distinct E); (1) with at least paired groups of setae
spines directly adjacent to the row. Dinoth- ranging from whitish-grey to golden brown
enarus badipes and S. caesareus lack an outer (same colour as other non-black setae; Fig. 1F–
742 D. J. CLARKE
J). In C. flavipennis (scored as state 1) most MALE GENITALIA (MALE, M)

fringe setae are whitish-grey.
48. Apex of sternite VIII in females: (0) straight; (1) 49. Apex of male tergite X: (0) narrowly rounded; (1)
deeply bisinuate. State (1) is an autapomorphy subtruncate to shallowly emarginate (Fig. 3Q);
of Ht. wakefieldi. (2) very deeply emarginate.
Figure 9. Scanning electron micrographs of Creophilus imitator (A, B), C. maxillosus villosus (C–F, L, M), C. oculatus
(J, K), Hadrotes crassus (G), Xanthopygus xanthopygus (H, I) and Staphylinus caesareus (N, O). A, vertex showing
micropunctate microsculpture and shallow depressions (ch4-2, circle); B, same, detail of micropunctate microsculpture
(ch5-2) and two shallow depressions each with associated pore and microseta (ch4-2, arrows); C, vertex showing small
abrupt pits (ch4-1, arrow); D, left procoxa and femur of large male with basal femoral spine (ch36-1, arrow), ventrolateral;
E, left protibia with mesal pair of protibial spines (ch37-1, arrows), posterior; F, prothorax with arcuate sternacoxal ridge
(ch23-1, upper arrow) and medially extended mesothoracic peritremes (ch25-1, lower arrow), ventral; G, prothorax with
anteromedially projecting sternacoxal ridge (ch23-0, arrow), ventral; H, prothorax showing anterior (left arrow), inferior
(middle arrow), and superior (right arrow) pronotal marginal lines, ventral-oblique; I, pterothorax showing two distinct
ridges of mesepimeron (ch27-0, arrows), lateral; J, prothorax showing anterior (left arrow), inferior (middle arrow), and
superior (right arrow) pronotal marginal lines, lateral-oblique; K, pterothorax showing single distinct ridge of
mesepimeron (ch27-1, arrow), lateral; L, left side of abdominal segment II showing position of one muricate fossa (ch41-1,
arrow), dorsal; M, surface detail of vaginal plate; N, female internal genitalia showing sclerotized plate at opening of
bursa copulatrix (ch87-1, left arrow), ciliose vaginal plate (ch85-0, middle arrow), and extensive muricate region (vaginal
fold) distal to vaginal plate (right arrow); O, surface detail of K. Scale bars: B = 0.01 mm; A, C, L–O = 0.1 mm;
F–K = 0.5 mm; D = 1 mm.
䉳
50. Apex of male sternite IX: (0) deeply and 58. Lateral inflexed teeth at apex of median orifice:
narrowly emarginate; (1) narrowly rounded; (0) absent (Figs 3 K, 31D, E); (1) present (Fig. 3,
(2) subtruncate to shallowly emarginate lit).
(Fig. 3Q). 59. Paramere: (0) fused to median lobe basally, not
51. Asymmetrical basal portion of male sternite IX: dorsoventrally movable; (1) not fused to median
(0) simple short process; (1) extended into elon- lobe basally, dorsoventrally movable.
gate process (Fig. 3Q). 60. Shape of paramere: (0) lanceolate, gradually
52. Ventral face of apical portion of median lobe: (0) narrowing apically (Fig. 3L, N); (1) spatulate,
continuously sclerotized (Fig. 3A); (1) divided gradually widening apically (e.g. Fig. 13C); (2)
longitudinally by membranous strip (Fig. 3C, widened basally, abruptly narrowed apically
arrow). (Fig. 3M).
53. *Apical subtriangular membranous region of 61. Condition of parameral apex: (0) narrowly
median lobe (ventral side): (0) absent (Fig. 3O); rounded to subtruncate (e.g. Figs 3L, N, 23C);
(1) present (Fig. 3J, stm). (1) notched (Fig. 3M, 13C).
54. *Relative length of median orifice: (0) elongate; 62. *Basal inner tube of internal sac: (0) absent;
(1) truncate or very short. This character (1) present (Figs 3, 4, bit).
describes length variation in the apical portion of 63. *Length of basal inner tube of internal sac
the median lobe, measured from the ventral-most relative to length of copulatory piece: (0)
medial part of the median orifice to the dorsal- shorter than straight portion of copulatory
most part. In C. erythrocephalus, C. lanio, C. piece (e.g. Fig. 3A); (1) much longer than
imitator, and C. albertisi this part of the median straight portion of copulatory piece.
lobe is distinctly ‘compressed’, such that the 64. *Basal cap of inner tube of internal sac: (0) absent
apical part seems truncate despite always having (e.g. Fig. 31E); (1) present (Figs 3, 4, bc).
a dorso-apical projection, whereas in all other 65. *Basomedial sclerotized strip of ventral side of
ingroup species it is elongated. internal sac when everted: (0) absent (e.g.
55. Dorsal–apical edge of median orifice: (0) not Fig. 3, C, J); (1) present (Fig. 3K, bms). This
carina-delimited, apex of median lobe not dis- distinctive sclerotized region of the internal sac
tinctly separate from internal part of median lobe is subcontinuous with the median lobe ventro-
(Fig. 3P); (1) carina-delimited, apex of median medially, and thus might also be considered an
lobe distinctly separate from internal part of extension of the ventral sclerotization of the
median lobe (Fig. 3O, ce). In C. imitator the median lobe.
carina is partly obsolete medially. 66. *Paired basoventral spiculose regions: (0)
56. *Paired apicolateral sclerites of median lobe: (0) absent (Fig. 3K); (1) present (Fig. 3A-D, H, sr).
absent (Fig. 3K); (1) present (Fig. 3, as). 67. *Ventromedian spiculose strip: (0) absent
57. *Attachment of apicolateral sclerites: (0) sepa- (Fig. 3K); (1) present (Fig. 3A-E, H, J, vr).
rated from lateral edges of median orifice 68. *Ventral process of internal sac: (0) absent
(Fig. 3A); (1) fused with lateral edges of median (Fig. 3K); (1) present, well developed (Fig. 3D,
orifice (Fig. 3J). E, vp); (2) present, reduced (Fig. 3H, vp).
744 D. J. CLARKE
Figure 10. Scanning electron micrographs of copulatory pieces. A, Creophilus flavipennis; B, C. maxillosus villosus; C,
C. erythrocephalus. Abbreviations as in Figures 3 and 4. Scale bars = 0.1 mm.
69. *Ventral sclerite of internal sac: (0) absent 76. *Dorsal receptacle for apex of copulatory piece/
(Fig. 3K); (1) present (Fig. 3A-E, H, J, vs). appendix: (0) absent; (1) present (Fig. 3O, P, dr).
70. *Double copulatory piece: (0) absent (Fig. 3K); 77. *Membranous lobe dorsal to the dorsal receptacle:
(1) present (Fig. 3A–E, H, cp, Fig. 4). (0) absent (Fig. 3P); (1) present (Fig. 3O, ml).
71. *Structure of double copulatory piece of inter- 78. *Styli of males and females: (0) without spine-like
nal sac: (0) straight, not cane-like (Fig. 3E, cp); bristles; (1) with dense covering of spine-like
(1) elongate, distinctly cane-like (Fig. 3A–D, H, bristles on ventral side. State (1) is an autapo-
J, cp). morphy of Hp. fossor.
72. *Parallel oblique ridges on dorsal side of copu-
latory piece: (0) absent (Figs 4A, D, G–O, 10C);
(1) present (Figs 4B, C, E, F, 10A, B, por).
FEMALE GENITALIA (FEMALE, F)
73. *Location of appendix in relation to apex of
copulatory piece: (0) extending beyond apex of 79. Apex of abdominal tergite X in female: (0)
copulatory piece (Figs 3F, 4A–F, N, 10A, B, broadly rounded (Fig. 23L); (1) subtruncate to
app) (1) enclosed by apex of copulatory piece shallowly emarginate.
(Figs 4G–K, 10C, app, iedg); (2) indistinguish- 80. First and second gonocoxites: (0) separated by
ably fused to copulatory piece (Fig. 4L, O, app). membranous suture (Fig. 5D, I, gc); (1) fused
The presence of an appendix in T. pictus could (Fig. 5J). This character is parsimony uninforma-
not be confirmed. tive. State (1) is an autapomorphy of T. pictus.
74. *Structure of appendix: (0) produced to point 81. Gonostyle of female: (0) articulates with second
(e.g. Figs 3F, 4E, F, 10A, app); (1) flared gonocoxite apically, separated by distinct mem-
(Figs 4C, D, 10B, app); (2) produced into membranous joint (Fig. 5I, arrow, sty); (1) articulates
branous swelling (shown as situated more or with second gonocoxite subapically (Fig. 5D, sty);
less completely enclosed by apical inner edges (2) fused to second gonocoxite, or absent (Fig. 5J).
of arms of copulatory piece in Fig. 4G, iedg). State (2) is an autapomorphy of T. pictus.
75. *‘Cross-bar’ of appendix: (0) absent (e.g. 82. *Vaginal fold: (0) membranous (Fig. 5A, vf); (1)
Fig. 4D); (1) present (Fig. 4B, C, E, F, xbar). with single broad sclerite (Fig. 5C, vf); (2) with
two smaller lateral sclerites (Fig. 5B, vf); (3) with (Fig. 8A, am); (1) anterior to, or in line with,
single narrow medial sclerite (Fig. 31I, vf, arrow). antennal insertion (Fig. 6, am).
83. *Median groove of vaginal plate/bursa copula- 93. Anterolateral macroseta of head (al): (0) absent;
trix: (0) absent (Fig. 5F, G); (1) present (1) present.
(Fig. 5A–C, mg). This groove continues from the 94. Basiantennal macroseta of head (ba): (0) absent
vaginal plate into the opening of the posterior or vestigial, and never a pair; (1) present, fully
part of the bursa copulatrix. developed.
84. *Median sclerotized strip of female vaginal plate: 95. Number of supraorbital (so), temporal (tm), and
(0) absent (Fig. 5B, C, F, G); (1) present (Fig. 4A, occipital (oc) macrosetae of head: (0) each single;
ss). (1) each with cluster of 2–4.
85. *Surface of female vaginal plate: (0) ciliose 96. Additional temporal macrosetae of head (ad): (0)
(Fig. 9N, middle arrow, O); (1) with minute absent; (1) present.
embedded teeth (Fig. 9M); (2) smooth. The con- 97. Genal macroseta of head (gn): (0) absent; (1)
struction of this character represents the hypoth- present.
esis that (0) and (1) are alternative forms of the 98. Position of temporal (tm) and occipital (oc) mac-
same homologous outgrowths of the vaginal plate rosetae of head: (0) anterior to posterior margin
surface. of head; (1) close to posterior margin of head
86. *Vaginal plate: (0) unsclerotized (Fig. 5A, vp) (Fig. 6A). This character is used to separate
(1) with pair of lateral sclerites (separated medi- groups of genera in Staphylinina (see Newton
ally) (Fig. 5B, C, pls); (2) with continuously scle- et al., 2000).
rotized ring (Figs 5F, 31I, vp); (3) completely 99. Ventroapical scapal macrosetae (vs): (0) absent,
sclerotized; (4) partially sclerotized only distally or strongly reduced; (1) two present.
(Fig. 21H, vp). State (4) is an autapomorphy of C. 100. Apical pairs of setae on antennomere 11: (0) both
flavipennis. pairs closely situated anterior to antennal
87. *Sclerotized plate at opening of bursa copulatrix: midline (e.g. Figs 25C, 27B); (1) pairs widely
(0) absent (e.g. Fig. 5H); (1) present (Fig. 5A, G, separated, one on either side of antennal
sc, Fig. 9N, left arrow, O). In Ht. wakefieldi and midline (e.g. Figs 13F, 16B, 23B).
D. badipes (scored state 1) this structure is
present but tiny in comparison with all other
species scored state (1). Thorax
88. Bursa copulatrix: (0) absent (Fig. 5H); (1) present 101. Number of macrosetae on anterior pronotal
(Fig. 5A–C, F, G, bcx). The bursa copulatrix is angle (anterior pronotal group, ap): (0) zero; (1)
considered absent if there is no pouch-like struc- cluster of more than two.
ture immediately anterior to the orifice that 102. Basolateral pronotal macroseta (bp): (0) absent;
connects the spermatheca to the vaginal cavity. (1) present.
89. *Spermathecal attachment: (0) to side of bursa 103. Basisternal macrosetae (bs): (0) absent; (1)
copulatrix (Fig. 5F, G, bcx, sp.); (1) to common present, as pair.
oviduct (Fig. 5H, sp.); (2) to base of flagellum 104. Parascutellar 1 of elytral parascutellar series
(Fig. 5A–C, fl, sp.). (ps1): (0) absent; (1) present.
90. *Flagellum of female bursa copulatrix: (0) absent 105. Number of macrosetae in elytral discal series
(Fig. 5F, H); (1) present (Fig. 5A–C, G, fl). (ds): (0) zero; (1) one or two; (2) three to eight.
91. *Structure of female spermatheca: (0) abruptly 106. Number of elytral basohumeral macrosetae (bh):
sacculate, with storage sac distinctly bulbous (0) zero; (1) one; (2) two.
(variable in shape) (Fig. 5A, F, H, sp); (1) more or 107. Number of macrosetae in posterior elytral patch
less threadlike in structure or gradually expand- (pt): (0) zero; (1) one; (2) three or more.
ing apically into storage sac (Fig. 5B, C, G, sp). 108. Metaventrital discal macroseta (mv): (0) absent;
(1) present.
109. Posteromedial metaventrital macrosetae (pm):
*ADULT CHAETOTAXY (CHAETOTAXY, C) (0) one pair; (1) two or more pairs.
110. Procoxal patch (pp): (0) absent; (1) present.
Unless otherwise indicated, all setae and abbrevia-
111. Number of macrosetae on distal edge of procoxa
tions listed below are illustrated in Figure 6.
(apical procoxal group, pc): (0) zero; (1) one; (2)
two; (3) more than two.
Head 112. Number of protrochanteral macrosetae (tr): (0)
92. Position of anteromarginal macroseta of head one; (1) two; (2) more than two.
(am): (0) posterior to antennal insertion 113. Mesocoxal patch (mp): (0) absent; (1) present.
746 D. J. CLARKE
114. Number of macrosetae on distal edge of meso- compatible with each other, much resolution is lost in
coxa (apical mesocoxal, mc): (0) one; (1) two; (2) the taxonomic congruence tree due to conflict between
more than two. each of the male genital and chaetotaxic character
115. Number of mesotrochanteral macrosetae (mt): (0) systems, and the other two: Hp. fossor and node 13
zero; (1) one; (2) more than two. are placed in a clade with Creophilus in the former,
116. Metacoxal patch (me): (0) absent; (1) present. whereas X. xanthopygus is placed within the ingroup
117. Number of metatrochanteral macrosetae (ma): in the latter. Those arrangements collapse most reso-
(0) zero; (1) one or more. lution in the taxonomic congruence tree.
According to the most basic criteria for assessing
dataset quality (see Table 1), the four character
Abdomen
systems studied here in general fall into the
118. Location (when present) of inner lateral (il) and
following sequence (from ‘best’ to ‘worst’): ‘structure’ >
median (mm) macrosetae of abdominal tergites
‘male’ > ‘female’ > ‘chaetotaxy’. Structural characters
III–VI: (0) close to middle of tergites (e.g. as in
generally performed better than the others, having
Fig. 6C, tergites VII and VIII); (1) at apex of
the most parsimony-informative characters, least
tergites (as in Fig. 6C, tergites III–VI).
homoplasy, fewest MPTs, most resolution, and the
119. Median macroseta (mm) of abdominal tergite III:
most nodes in common with the simultaneous analy-
(0) absent; (1) present.
sis tree (Table 1; Fig. 11). By these same criteria,
120. Median macroseta (mm) of abdominal tergite IV:
chaetotaxy performed the worst, and in large part
had values opposite to or much worse than the
121. Median macroseta (mm) of abdominal tergite V:
structural characters, the most notable being the
extremely large number of MPTs and the low CI
122. Median macroseta (mm) of abdominal tergite VI:
(0.45). The length and fit of the structural characters
is nearly the same on the MPTs for both the simul-
123. Median macroseta (mm) of abdominal tergite
taneous and the separate analyses (reflecting the
VII: (0) absent; (1) present.
nearly identical topologies), whereas each of the other
124. Inner lateral macroseta (il) of abdominal tergite
three character systems had increased length and
III: (0) absent; (1) present.
more homoplasy on the simultaneous analysis MPTs
(Table 1). Thus, both topology and tree statistics dem-
IV: (0) absent; (1) present.
onstrate the dominant influence of external structural
characters in the simultaneous analysis, despite com-
V: (0) absent; (1) present.
prising only about one-third of the total number of
characters.
VI: (0) absent; (1) present.
Nevertheless, the other three character systems
positively influenced the simultaneous analysis in
VII: (0) absent; (1) present.
several ways. First, although resolution is particu-
larly poor for female genitalia and chaetotaxy
VIII: (0) absent; (1) present.
in separate analyses, the number of parsimony-
130. Parascleral macroseta (pa) on outer parasclerites
informative female characters/states (12/32) is
of abdominal segments III–VI: (0) absent; (1)
comparatively small (Table 1: PIC; Table 2: no. of
present. State (1) is an autapomorphy of Ht.
character states), and in separate analyses of both
crassus.
character systems a high proportion of recovered
nodes (80 and 63%; Table 1) is congruent with the
simultaneous analysis tree. This demonstrates that
RESULTS OF PHYLOGENETIC ANALYSES
even apparently poor character systems (few charac-
PHYLOGENETIC UTILITY OF CHARACTER SYSTEMS ters, low CI) are contributing some important phylo-
Two features of the separate analyses stand out genetic signal to an analysis of ‘total’ morphology.
(Fig. 11). First, external structural characters clearly Second, Figure 12A demonstrates that in terms of
dominate the analysis: the consensus tree for these synapomorphy, non-structural systems are crucial for
characters is nearly identical to the combined analy- providing the strongest hypothesis of relationships:
sis tree. Second, the taxonomic congruence tree is some nodes are not supported by structural synapo-
completely unresolved, thus demonstrating a lack of morphies (nodes 8 and 17), other nodes are supported
compatible components among the four character primarily by other character systems (e.g. nodes 4, 5,
systems. Close inspection of the separate consensus 7, and 16), and others have comparable character
trees reveals, however, that although the trees for support from more than one system (nodes 7, 10, 13,
female genitalia and structural characters are largely and 14). In fact, the proportion of non-structural
Figure 11. Strict consensus trees from analysis of combined and separate character systems. Grey dots indicate nodes
congruent with the analysis of all the data. Numbers above branches indicate Bremer support; numbers below branches
are numbered nodes referred to in the text. The taxonomic congruence (Mickevich, 1978) tree represents the strict
consensus of trees obtained in the separate analyses. Tree statistics are presented in Table 1.
Table 1. Tree statistics for separate and combined phylogenetic analyses
Data No. of Total no. No. of congruent

partition PIC (%) MPTs TL *TL-sim. CI *CI-sim. RI *RI-sim. of nodes nodes (%)
S 44 (36) 3 91 91.82 0.71 0.71 0.89 0.89 17 16 (94)

M 29 (24) 16 55 59.82 0.64 0.59 0.86 0.83 13 10 (77)
F 12 (10) 182 30 36.0 0.67 0.56 0.86 0.77 5 4 (80)
C 36 (30) 3551 96 109.36 0.45 0.39 0.68 0.60 8 5 (63)
All data 121 11 297 – 0.55 – 0.78 – 18 –
PIC, parsimony-informative characters; No. of MPTs, number of most-parsimonious trees; TL, tree length; *TL-sim.,
average tree length on simultaneous analysis MPTs; CI, consistency index; *CI-sim., average consistency index on
simultaneous analysis MPTs; RI, retention index; *RI-sim., average retention index on simultaneous analysis MPTs; S,
external structures; M, male genitalia, F, female genitalia; C, chaetotaxy.
character state changes is equal to or greater than therefore reveal that non-structural characters, when
the proportion of structural character state changes considered together, are at least as important for
for 15 of the 18 (~83%) nodes recovered in the simul- phylogenetic inference as the more easily studied
taneous analysis. The above features of the analysis external structural characters, and the result of
748 D. J. CLARKE
including them here is a much better supported
PBS, partitioned Bremer support (the total of PBS for all character systems = total BS); BS-sep, Bremer support in separate analysis; HPBS, hidden partitioned
Bremer support; min. steps, minimum number of steps required for a character system; PSYN, partitioned unambiguous synapomorphy; SYN-sep, unambiguous
synapomorphies in separate analysis; HPSYN, hidden partitioned unambiguous synapomorphy; PUNIQ, partitioned uniquely optimized synapomorphy;
UNIQ-sep, uniquely optimized synapomorphy in separate analysis; HPUNIQ, hidden partitioned uniquely optimized synapomorphy; S, external structures; M,
SPUNIQ/
partition SPBS SBS-sep SHPBS steps min. steps SPSYN SSYN-sep SHPSYN char. states of char. states SPUNIQ SUNIQ-sep SHPUNIQ SPSYN
tree.
male genitalia, F, female genitalia; C, chaetotaxy. The summation sign (S) indicates that each cell is the sum of values across 18 nodes in Figures 11–12.
0.57
0.69
0.52
0.70
0.41
Structural characters also dominated measures of
overall support in the simultaneous analysis, contrib-
Uniquely optimized synapomorphy
uting 52% of the total PBS, 42% of the total PSYN,

and 50% of the total PUNIQ (Table 2). However, a
substantial proportion (~28%) of PBS is derived from
-2
2
4
4
8
male genitalia, and this character system is most
comparable with chaetotaxy in numbers of PSYN and
PUNIQ. The contribution of female genital characters
to absolute support is clearly negligible, at only ~2%
of the total BS, and ⱕ 10% of the total PSYN and
37
13
3
9
62
PUNIQ (Table 2). In some cases, correcting for differ-
(UNIQ)
ences in partition size changed the relative contribu-

tions to overall support (Table 2; right column of each
35
15
7
13
70
group): support contributed by structural and male

genital characters is very similar for PBS and PSYN,
SPSYN/ no.
while structural characters are clearly superior to

male genitalia in absolute support. Relative values of
PSYN and PUNIQ are much more comparable among
0.43
0.47
0.45
0.31
0.41
character systems and relative female PUNIQ is even

slightly greater than that of structural characters
(0.70 vs. 0.68; Table 2: PUNIQ/PSYN). The number of
female PUNIQ relative to PSYN therefore suggests
No. of
that some female genital characters are strong

Table 2. Total support measures for each character system in the simultaneous analysis
109
64
32
79
284
hypotheses of primary homology.

All clades in the combined analysis are present in at
least one of the partition consensus trees (Fig. 11). This
is directly related to the strongly congruent phyloge-
-1
3
5
14
21
netic signal in the structural and male genital charac-

ters, which when analysed together recover 17 of the
18 nodes of the simultaneous analysis (results not
Synapomorphy
shown). Nevertheless, two major features of the

52
26
5
18
101
support analyses are 1) the high proportion of total

hidden Bremer support and 2) the overall high propor-
(SYN)
tion of hidden support (HPBS, HPSYN, and HPUNIQ)

51
29
10
32
122
contributed by chaetotaxy. Approximately 74% of the

total BS (= SPBS) was HPBS, and this contrasts
strongly with the much smaller proportion of HPSYN
min. SPBS/
and HPUNIQ (Table 2). Chaetotaxy contributed by far

0.59
0.76
0.77
0.40
0.11
the greatest amount of hidden support: 62% of HPBS,

66% of HPSYN, and 50% of HPUNIQ (Table 2). Sixteen
65
35
20
43
163
of the 18 nodes in the simultaneous analysis have

positive hidden Bremer support (supporting Table S1,
HBS). However, six of these nodes (2, 8, 11, 14, 15, and
70.99
4.69
12.94
9.25
44.11
17) are characterized by zero or negative hidden syna-

pomorphy (supporting Table S2, HSYN). Examination
of the different character systems for a discrepancy
between HPBS and HPSYN reveals an important
49.69 45
26.94 14
2.25 -7
17.11 -27
95.99 25
contrast between non-chaetotaxy and chaetotaxy

characters (supporting Table S1, HPBS; Table S2,
Bremer support
HPSYN): structural characters contributed positive

HPBS to five nodes, but these nodes have zero HPSYN;
All data
male genitalia contributed positive HPBS to 11 nodes,

Data
but eight of these (73%) have zero HPSYN and female

(BS)
genitalia contributed positive HPBS to six nodes, but

C
F
S
Staphylinus caesareus
3/2/-2/0 Platydracus maculosus
81 Ontholestes cingulatus
11 Dinothenarus badipes
2/1/-1/0
Xanthopygus xanthopygus
61 2.5/1.25/-0.75/0 Hadrotes wakefieldi
1 66 6.33/-0.67/-2/5.33 Thinopinus pictus
5/4/1/1
2 92 Hadropinus fossor
99
12 1/2/0/1 Hadrotes crassus
(A) 3 1/6/0/0
84 3/2/0/2 Liusus hilleri
94
13 100 Liusus humeralis
Creophilus complex 4 14 C. variegatus
2.5/2/0/2.5 5.18/-1.82/0/1.64 C. m. maxillosus
94 97 C. m. villosus
0.18/0.18/2/-0.36
structure 5 15 C. incanus
male 82
female 8/0/1/0 16 C. flavipennis

C. galapagensis
chaetotaxy 99
C. oculatus
6 0/3/0/2
2/0/0/1
99 C. huttoni
4/4/3/1
17 96 C. rekohuensis
Creophilus 100 18
14 28 50 51 90 91 97
7 0/2/
0/-1
C. lanio
0 1 1 1 1 1 0 Staphylinus caesareus 1/0/
* 58 85 86 87 88 89 106 1/0 C. erythrocephalus
16 54 59 60 127 Platydracus maculosus 65
1 2 3 0 0 1 2
3/0/0/1
1 1 1 1 1
+
10 70 112
2 1 1 Ontholestes cingulatus 8 85 C. albertisi
11 351 901 1062 Dinothenarus badipes 9 91 C. imitator
15 26 52 128 * 5 23 26 105 107 123 129
Xanthopygus xanthopygus
10
3 1 2 0 0 0 0
0 1 1 0 +++ 5 48 55 86 88 128
Hadrotes wakefieldi ** * * + ++
1 22 44 45 56 101 121122
1 1 1 1 0 0 0
2 1 1 3 0 1
5 11 12 14 35 38 43 87 101102105 109112113114116
2 7 24 30 32 41 42 55 64 67 69 80 81 86 90 93 103 107 111115 117 123124125126127
2 0 0 0 1 0 1 1 0 0 0 1 2 2 0 0 0 0 0 2 1 0 0 0 0 0 Thinopinus pictus
* +
2 4 7 19 24 27 40 50 51 61 65 79 99 106 110 124 127 1 1 1 0 2 2 1 0 1 0 0 1 1 1 2 1 78 94 95 97 111
+ Hadropinus fossor
12
3 1 0 1 1 1 2 1 1 0 1 0 0 1 1 1 1 0 1 0 3
+ 16 35 94 97 104105107109114128130
Hadrotes crassus
3 15 28 45 58 59 62 64 67 68 69 70 76 90
30 38 46 63 73 95 111
1 1 1 1 2 1 2
0 2 0 0 0 0 2 1 1 1 1
*5 29 31 52 *53 ++ Liusus hilleri
96 110
1 1 0 1 1 1 1 1 1 1 1 1 1
* ** * * 13 1 1 1 0 1 1 0 57
Liusus humeralis
14
1
(B) Creophilus complex 4 40 55 57 61 66 79 94 103 128

0 1 1 0 0 0 0 0 1 C. variegatus
16 39 71 83 104106119120121122 4 20 21 36 47 100 111
* C. m. maxillosus
1 1 1 1 1 2 1 1 1 1 1 1 1 1 1 1 0 74
* * * + C. m. villosus
15
1
5 17 84 91 114 127
1 1 0 1 0
10 102 122
1 0 0 C. incanus
+ 86 111 127
8 9 10 13 23 25 26 37 66 81 105 107 16 4 2 1 C. flavipennis

34 72 74 75 103104106119120121122123124125126129
1 3 1 1 1 1 0 1 1 1 2 2 0 0 1 0 0 0 01 0 0 0 0 0 0 0 0 0 C. galapagensis
* +
6 53 57 74 112 115
C. oculatus
1 1 2 1 1 26 35 105 C. huttoni
1 6 18 45 52 68 73 77 82 85 86 106108119 17 2 1 1 10
C. rekohuensis
Creophilus 1 1 1 1 0 2 1 1 1 2 1 1 1 0
**
18 0
7 104 108
C. lanio
0 0
54 55
1 1
* C. erythrocephalus
15 82
2 58 59 64 77 82 83 86 104108 122
8 0 2 4 5 20 94 127 129 3 0 0 0 0 3 0 2 0 0 0 C. albertisi
9 2 2 2 0 0 0
+ 102 119 123 *
C. imitator
10
0 1 0
Figure 12. Strict consensus of 11 most-parsimonious trees from the combined analysis of 121 adult morphological
characters of the Creophilus complex (TL = 297; CI = 0.55; RI = 0.78). A, bars indicate the proportional contribution of each
character system to the total synapomorphies supporting each clade. Numbers above branches are partitioned Bremer
support values for structure, male genitalia, female genitalia, and chaetotaxy characters, respectively; small numbers
below branches are bootstrap proportions. B, filled squares = uniquely optimized character state changes, open
squares = homoplasious changes. Character and state numbers are given above and below branches, respectively.
* = ACCTRAN; + = DELTRAN. Taxa in bold type are marine littoral species. Large numbers 1–18 indicate nodes referred
to in the text.
750 D. J. CLARKE
three of these (50%) have zero HPSYN. Chaetotaxy, in monophyletic (node 12: BP = 92%; BS = 9) and it is
contrast, contributed positive HPBS to the majority evident from Figure 12B that these species are mor-
of nodes (13 of 18), and most of these (10 of 13; 77%) phologically highly derived. This clade is the sister-
are characterized by positive HPSYN. There are few group of the remaining genera (node 5), which are
hidden uniquely optimized synapomorphies in the divided into two very strongly supported monophyl-
simultaneous analysis, and many of the PUNIQ etic groups. Node 13 comprises Hadrotes crassus
contributed by female genitalia and chaetotaxy (Hadrotes sensu stricto) + Liusus (BP = 84%; BS = 4).
are recovered only in the simultaneous analysis as The monophyly of Liusus (node 14: BP = 100%;
hidden unique synapomorphies (supporting Table S3, BS = 7), the only other non-monotypic genus, is
HPUNIQ). strongly supported by seven synapomorphies (three
structural, two male genital, two chaetotaxic), three
of which are unique: intercoxal tumescence of
SIMULTANEOUS ANALYSIS OF ALL ADULT DATA metaventrite present (ch29-1); central part of
The simultaneous analysis of all 121 characters pro- metaventrital disc glabrous (ch31-1); additional tem-
duced 11 shortest trees. Statistics for the analysis are poral macrosetae of head present (ch96-1). Node 6
presented in Table 1. The strict consensus tree comprises the genus Creophilus (BP = 99%; BS = 9),
(TL = 299) with mapped character state changes is monophyly of which is supported by 12 synapomor-
shown in Figure 12B, and unique (single origin) phies (eight structural, one male genital, one female
changes are summarized below for clades of greatest genital, two chaetotaxic), nine of which are unique:
interest to the analysis. Most nodes recovered in antennae clavate, with antennomeres 7–10 gradu-
the simultaneous analysis tree are very strongly sup- ally increasing in diameter (ch8-1); antennal micro-
ported. Of 18 nodes, 11 have BP > 90% and nine have pubescence only on antennomeres 7–11 (ch9-3);
BS ⱖ 5 (Figs 11, 12). Ambiguity among the 11 solu- ventral basolateral ridge of left mandible present
tions resulted from alternative resolutions of a poly- (ch13-1); mesothoracic spiracular peritremes very
tomy within the maxillosus-group (node 16). large, expanded medially with posterior edge pro-
Figure 12 shows several systematically important duced into point (ch25-1); mesoventral intermeso-
results, the first being the monophyly of the Creophi- coxal process truncate or broadly rounded (ch26-0);
lus complex (node 3: BP = 99; BS = 11). Of the 16 pair of spines on inner edge of protibia present
synapomorphies supporting this group (six structural, (ch37-1); paired basoventral spiculose regions of
four male genital, one female genital, five chaetotaxic) internal sac present (ch66-1; ACCTRAN optimiza-
13 are uniquely optimized: punctation of vertex tion); gonostyle of female articulates with second
absent (ch4-3); ventral basal ridge separate from gonocoxite subapically, the point of articulation not
ventral portion of postoccipital suture near basal visible in ventral view (ch81-1); three to eight mac-
gular impression (ch7-1); vestiture of head and prono- rosetae in elytral discal series (ch105-2).
tum absent (ch19-0); posteromedian furcasternal Fourth, the monophyly of both the maxillosus- (node
fovea present (ch24-1); mesepimeron with single ridge 15) and erythrocephalus- (node 7) groups is very
at edge of sclerite (ch27-1); anterolateral flanges of strongly supported, so Newton’s (1985) informal
abdominal tergite II poorly developed (ch40-1); male species groups are here corroborated. Monophyly of the
sternite IX subtruncate to shallowly emarginate maxillosus-group (BP = 97%; BS = 5) is supported by
(ch50-2); parameral apex notched (ch61-1); basome- seven synapomorphies (five structural, two chaeto-
dial sclerotized strip of internal sac absent (ch65-0); taxic), six of which are unique: vertex with small
apex of abdominal tergite X in females subtruncate to abrupt pits (ch4-1); basolateral pronotal impressions
shallowly emarginate (ch79-1); ventroapical scapal asperate and setose (ch20-1); largest males with baso-
setae absent or strongly reduced (ch99-0); patch of lateral pronotal denticle (ch21-1); large males with
macrosetae on procoxa present (ch110-1); inner basal spine-like projection on profemur (ch36-1); apical
lateral macrosetae of abdominal tergite III present fringe setae of abdominal segments III–VI with paired
(ch124-1). Second, Ht. wakefieldi is placed as the groups of whitish-grey or golden brown setae (ch47-1);
sister-species of all other Creophilus complex species, apical pairs of setae on antennomere 11 widely sepa-
supporting earlier suspicions that Ht. wakefieldi was rated, one on each side of antennal midline (ch100-1).
misplaced in Hadrotes (Newton, 1973; Klimaszewski Of particular note within this group is the lack of
et al., 1996), yet supporting a relationship with the resolution within a very strongly supported subclade
rest of the genera. As Hadrotes is therefore polyphyl- (node 16: C. m. maxillosus, C. m. villosus, C. incanus,
etic as currently delimited, Ht. wakefieldi should be C. flavipennis, and C. galapagensis). This polytomy is
placed into a different genus. a result of some character conflict but also, more
Third, the two monotypic genera Hadropinus and importantly, a general lack of phylogenetically infor-
Thinopinus are together strongly supported as a mative characters among maxillosus-group species.
Most intriguing here is the lack of support for a within Staphylinidae to compare ‘conventional’ exter-
monophyletic C. maxillosus (and C. m. villosus and C. nal structural characters with a broad sample of
m. maxillosus together were not recovered as a clade in largely ‘unconventional’ characters. The motivation
any of the 11 optimal solutions). for this study was therefore the premise that survey-
Monophyly of the erythrocephalus-group (node 7: ing multiple character systems for a finite taxon
BP = 100%; BS = 12) is supported by 14 synapomor- sample should produce the best estimate of phylogeny
phies (four structural, four male genital, three female because increasing the number of characters should
genital, three chaetotaxy), nine of which are unique: increase phylogenetic accuracy (Wiens, 2004, and ref-
head at least partly orange or yellowish (ch1-1); heads erences therein).
of large males with ventrolateral carina (ch6-1); man-
dibular tooth T3 proximal to mandibular blade (ch18- PHYLOGENETIC SIGNAL IN DIFFERENT
1); dorsal process of internal sac reduced (ch68-2); CHARACTER SYSTEMS
appendix of internal sac enclosed by apex of copula-
External structural characters contributed the stron-
tory piece (ch73-1); membranous lobe situated dorsal
gest phylogenetic signal. Analysed separately, they
to dorsal receptacle for copulatory piece/appendix
produced the most-resolved and best-supported con-
present (ch77-1; reversed in C. albertisi); vaginal fold
sensus tree and clearly had the greatest influence on
with single broad sclerite (ch82-1; further changed
the topology of the combined analysis (Fig. 11). But
above); vaginal plate with pair of medially separated
separate analyses of the other character systems
sclerites (ch86-1; further changed in C. albertisi);
revealed that all four contributed distinct phyloge-
metaventrital discal macrosetae present (ch108-1;
netic signal that was largely a subset of that in the
ACCTRAN optimization). In contrast to the
simultaneous analysis (Fig. 11; Table 1, no. of congru-
maxillosus-group, phylogenetic relationships within
ent nodes). The overall effect of combining these data
the erythrocephalus-group are completely resolved
was therefore to amplify the signal recovered by the
and moderately to strongly supported. Basally, the
structural characters (Fig. 12A).
erythrocephalus-group is split into a New Zealand
But simultaneously analysing these characters also
clade (node 17: BP = 99%; BS = 5) and an Australa-
had the effect of introducing some complementary
sian clade (node 8: BP = 65%; BS = 1). Within the
phylogenetic signal and thus improving resolution in
latter, C. imitator and C. albertisi (node 10) form a
parts of the tree where individual character systems
strongly supported group endemic to New Guinea
were either lacking or in conflict. For example,
(BP = 91%; BS = 4).
there were no female genital characters that
could support monophyly of Liusus (node 14),
DISCUSSION AND CONCLUSIONS Hadropinus + Thinopinus (node 12), or the New
Zealand Creophilus species (node 17), so the female
A general goal of systematics is to understand the
consensus tree is under-resolved (Fig. 11). But the
utility of different classes of data so that we can
poorly resolved chaetotaxy consensus tree, due mostly
establish robust phylogenies and classifications. But
to character conflict, nevertheless recovered all of
because of a preoccupation with comparing morpho-
those clades. Similarly, the New Zealand Creophilus
logical with molecular characters, few studies have
clade (node 17) is supported by male genital and
compared the phylogenetic utility of alternative
chaetotaxic, but not structural, characters, and only
organismal character systems – an especially impor-
chaetotaxy both supports this group and resolves
tant issue in groups where molecular data are
relationships within it. Localized phylogenetic signal
unavailable. Some papers, like this one, have focused
is widely held as a virtue of simultaneous analysis,
on the utility of characters traditionally perceived
especially in molecular phylogenetics, where, for
to be ‘unconventional’ or of low value (e.g. de Queiroz
example, fast-evolving mitochondrial and slow-
& Wimberger, 1993; Quicke & Belshaw, 1999;
evolving nuclear genes can resolve different levels of
Schander & Sundberg, 2001; Wiens, 2001; Strong,
the tree (e.g. Simon et al., 1994). Significantly, these
2003; Gaubert et al., 2005; Hill, 2005; Price, Fried-
results also indicate that chaetotaxy may be useful for
man & Omland, 2007; Lee, 2009; Meier & Lim, 2009;
resolving shallow nodes. Overall, the results are
Song & Bucheli, 2010), and a surprisingly frequent
encouraging in that they indicate much more agree-
conclusion is that such characters add significant
ment than disagreement.
information to an analysis and often show similar
levels of homoplasy to ‘conventional’ characters,
including DNA. For insects, the relative phylogenetic IF GENITAL STRUCTURES ARE UNDER SEXUAL
utility of alternative adult taxonomic character SELECTION, ARE THEY PHYLOGENETICALLY USEFUL?
systems is largely unknown (but see Song & Bucheli, As is typical for insects (Eberhard, 1985), male
2010 for recent example), and this study is the first genitalia are of major systematic importance in
752 D. J. CLARKE
Staphylinidae, and phylogenetic studies at or below Eberhard, 1985; Song & Bucheli, 2010), many of these
the tribal level (thus comparable with this study) studies would lack female genitalia entirely. The
usually include many male genital characters. For apparently unconventional use of female genitalia in
example, most studies of closely related species, staphylinid phylogenetics contrasts with some other
including most infratribal and infrageneric Staphyli- groups, where they (sensu lato) have sometimes been
nine group phylogenies, use ~ 25% male genital char- extensively employed, or have even formed the basis
acters (e.g. Herman, 1977, 1986, 1991; Thayer, 1997; for an entire analysis (e.g., Liebherr & Will, 1998:
Ashe & Chatzimanolis, 2003; Chatzimanolis, 2005, Carabidae). Like male genitalia, in some groups they
2008; Chatzimanolis & Ashe, 2005; Solodovnikov, may also be under sexual selection (via antagonistic
2006). But in some studies they are dominant: Sco- sexual coevolution: Eberhard, 1985). Probably the
paeina (Frisch et al., 2002: ~ 70% of 72 characters), best evidence available that this may be occurring in
Dolicaonina (Herman, 1981: 51% of 55 characters), the Creophilus complex is the almost equivalent taxo-
Pseudopsis (Herman, 1975: ~ 48% of 35 characters), nomic usefulness of female genitalia, a possibly
and Maorothius (Assing, 2000: 46% of 39 characters). unusual result (Eberhard, 1985). I am aware of no
Although these studies exemplify the use of male meta-analysis comparable with that of Song &
genitalia in phylogenetics, internal sac structures are Bucheli (2010), but Soulier-Perkins (2001) showed
not so widely used (see Thayer, 1985, for omaliine that in lophopid planthoppers female genitalia were
group example). In this study, > 50% of the 29 male critical for resolving both basal and terminal nodes,
genital characters were derived from the highly thus complementing the support for intervening
complex internal sac, and most of these provide nodes provided by male genitalia. My analysis is more
uniquely optimized synapomorphies. Male genitalia typical for Staphylinidae in including around 10%
are generally understood to be under sexual selection, female genital characters, but the fewer characters
and are thought to evolve rapidly. Because of this were of demonstrably high value (Table 2, PUNIQ/
male genitalia are often assumed to lack phylogenetic PSYN), and informative throughout the tree (Figs 11,
signal (see Song & Bucheli, 2010). But in a broad 12). Like male genitalia, resolution in separate analy-
survey of insect phylogenies, Song & Bucheli (2010) sis is also congruent with external structures and the
demonstrated that the phylogenetic utility of male combined analysis (Fig. 11). Overall, my analysis
genitalia is comparable with (and in many cases strengthens Song & Bucheli’s (2010) argument that
showing less homoplasy than) non-genital characters, sexual selection, if it is occurring, does not preclude
and consequently that male genitalia are indeed phy- phylogenetic utility – which must be determined in
logenetically useful regardless of mode and rate of the context of a simultaneous analysis.
evolution. The phylogenetic utility of male genitalia
in the Creophilus complex is highlighted by phyloge-
netic signal that is (1) largely congruent with that of PHYLOGENETIC UTILITY OF CHAETOTAXY – HIDDEN
structural characters (presumably not under sexual SUPPORT OR DISPERSION OF HOMOPLASY?
selection), and the simultaneous analysis, and (2) Within Staphylinidae, the use of chaetotaxy in phy-
spread throughout the phylogeny (Fig. 11) rather logenetics is inconsistent among groups and across
than localized to terminal nodes (an expectation of taxonomic ranks. The use of chaetotaxy as a phylo-
rapidly evolving characters). My results therefore genetic character system is widespread, with nearly
support their conclusions, and offer the first compara- all studies I have surveyed including at least some
tive empirical evidence for the phylogenetic utility of chaetotaxic characters. There is also a surprising
male genitalia in Staphylinidae. variability in the representation of chaetotaxic char-
Also typical for insects (Eberhard, 1985), female acters in published matrices. Some lack any (e.g.
genitalia are seemingly of less systematic importance Frisch et al., 2002), and most have few to moderate
in Staphylinidae, and are nearly always very poorly numbers of setal characters (most non-Tachyporine
represented in rove beetle phylogenies. The most group studies include less than 30% chaetotaxic char-
characters so far included were in a phylogenetic acters, e.g. Herman, 1970; Nomura, 1991; Leschen &
analysis of Pseudopsis (Herman, 1975: 20% of 35 Löbl, 1995; Smetana, 1995; Assing, 2000). Those with
characters). Female genitalia are absent from most a majority of chaetotaxic characters come from
infrageneric studies (e.g. all aforementioned infrage- studies of Tachyporine group subfamilies (> 40% of all
neric Staphylinine group studies; Wu & Zhou, 2007), characters in Kistner & Abdel-Galil, 1986 and Danoff-
but when included account for much less than 10% of Burg, 1994, with most others ~ 30%) and larval
the data (e.g. Smetana, 1995; Ashe, 2000, 2005; Ahn studies. Although chaetotaxy is used more in lower-
& Ashe, 2004), and are almost always characters of level analyses, Thayer’s (2000) analysis of the Oma-
the gonocoxites (external) or spermatheca (internal). liine group subfamilies (~ 90% larval chaetotaxic
As such, under some definitions of ‘genitalia’ (e.g. characters) demonstrated their application in even
higher-level studies. Ashe (1984) utilized 34% chaeto- tant to question whether this reflects the augmenta-
taxic characters in a study of adult gyrophaenine tion of additive signals or the cancellation of
aleocharines (Tachyporine group), and in a later inde- directional homoplasy in conflicting datasets (disper-
pendent larval study used 65% (Ashe, 1986). The sion of homoplasy: Barrett, Donoghue & Sober, 1991;
literature therefore suggests that in some groups Gatesy et al., 1999: fig. 3f). Distinguishing between
chaetotaxy provides many characters, even in adult these generally opposite effects (synergism vs. cancel-
analyses, but in others is very poorly represented. At lation) is necessary because synergism may indicate
present, we neither understand their relative phylo- phylogenetic accuracy whereas cancellation may
genetic utility, nor have any explanation for the simply be an analytical artefact. When hidden
inconsistent use of chaetotaxy within Staphylinidae. Bremer support results from dispersed homoplasy
In the Creophilus complex, adult chaetotaxy is com- there should be no hidden synapomorphies (Gatesy
parable with male genitalia as a source of characters et al., 1999: 280). Hence, a key result to emerge from
(Table 1). Yet this is more typical of Tachyporine the present study is the contrast between the hidden
group than other Staphylinine group studies, where support for chaetotaxy and the other character
chaetotaxy is relatively neglected. The dominance of systems. Most nodes with positive HPBS contributed
chaetotaxy in some analyses may simply reflect a lack by chaetotaxy had positive HPSYN contributed by
of other data, such as in larval analyses where adult- chaetotaxy, suggesting that HPBS contributed by cha-
specific characters are absent (genitalia, wings, etc.), etotaxy reflects synergistic signal amplification. In
but it is unlikely that Tachyporine group taxa are contrast, most nodes with positive HPBS contributed
richer, and adults of other groups poorer, in chaeto- by other characters did not have positive HPSYN,
taxic variation. So if chaetotaxy is potentially suggesting that most of that HPBS reflects dispersion
underutilized in staphylinid phylogenetics, the ques- of (some) homoplasy.
tion remains whether it is worthwhile developing this But another possibility belies this optimistic inter-
character system in other groups, especially given the pretation: combining a noisy subset of characters with
difficulty of studying chaetotaxy (Solodovnikov, 2007), a highly structured dataset could be expected to over-
and an anecdotal perception of lower value compared inflate both the character and Bremer support for
with more complex structural or genital characters. some well-supported clades by forcing the optimiza-
The following results could be seen as justification tion of many unambiguous (yet homoplastic, i.e.
for excluding chaetotaxy: (1) chaetotaxy contributed CI < 1.0) synapomorphies. Thus, a noisy character
the most homoplasy in both separate and combined system should be characterized by a lower ratio of
analyses (Table 1), and the fewest uniquely opti- HPUNIQ to HPSYN. A high percentage of the
mized synapomorphies (Table 2); (2) for ~ 64% of HPSYN contributed by male and female genitalia
the chaetotaxic characters the derived state(s) is are in fact HPUNIQ (Tables 2, S2, S3), indicating
‘absent’, and many absences are found in terminal that hidden support at nodes with positive HPSYN
taxa (e.g. C. galapagensis and T. pictus), suggesting reflects common signal amplification. For chaetotaxy,
that loss of macrosetae is common near the tips of a however, of the ten nodes with positive HPSYN, only
phylogeny, potentially introducing noise into the five of these had HPUNIQ (actual total HPUNIQ for
analysis; and (3) chaetotaxy has the greatest number these nodes = 7: supporting Table S3; the total listed
of polymorphic scorings (Appendix 1), reflecting more in Table 2 records net HPUNIQ for the dataset over
intraspecific variability than other characters, and all nodes), with the HPSYN of the other five nodes
again indicating evolutionary lability. But these representing homoplasies.
potentially negative features do not necessarily The existence of hidden support is often reported as
negate phylogenetic utility, and chaetotaxy provided a major result of including apparently unpromising
and enhanced resolution near the tips of the phylog- character systems in a simultaneous analysis.
eny and augmented the phylogenetic signal of other However, most studies conclude this without incorpo-
characters (Fig. 12A). Moreover, there is also chaeto- rating HSYN and HUNIQ as means of identifying
taxic evolutionary novelty that was recovered only in dispersed homoplasy (reminiscent of an increasing
simultaneous analysis, e.g. the metaventrital mac- trend of ‘phylogenies without synapomorphies’: Mooi
roseta (ch108-1) is a uniquely derived synapomorphy & Gill, 2010). My use of these metrics has enabled a
of the C. erythrocephalus group (Fig. 12B, node 7, more conservative interpretation of hidden chaetot-
ACCTRAN). axy support, and indicates that although chaetotaxy
Hidden signal/support (Chippindale & Wiens, 1994; contributed the most hidden Bremer support this
Nixon & Carpenter, 1996; Gatesy et al., 1999) is a does not necessarily represent positive signal ampli-
widely appreciated advantage of simultaneous analy- fication across the phylogeny. Despite the possible
sis. Given that much of the hidden Bremer support in advantages suggested by the hidden support analysis,
this study was contributed by chaetotaxy, it is impor- I therefore caution that the inclusion of chaetotaxy
754 D. J. CLARKE
must be weighed against the polymorphisms, coding suggests two equally parsimonious alternatives for
difficulties, and other issues discussed above. In any the evolution of the marine littoral association
case, my results are not overall condemning of cha- (Fig. 12) involving either two (at node 4 and in C.
etotaxy, and suggest that continued research into the huttoni) or three independent transitions (at nodes
phylogenetic utility of chaetotaxy is justified. 12, 13, and in C. huttoni), each requiring three
steps. The second scenario is preferred due to the
implausibility of a back-transition to the terrestrial
SYSTEMATICS OF THE CREOPHILUS COMPLEX environment (at node 6) The strong support for
Newton (1973) mentioned shared male genital char- nodes 12 and 13 in Figure 12 and the preference for
acters (internal sac structures) as indicating a close the second scenario argue for synonymizing the
relationship between all five genera in the Creophilus genera within each of these clades, thus removing
complex (but excluding Ht. wakefieldi). These obser- monotypic genera. But the mutual distinctiveness of
vations are corroborated here, with nine male genital Hp. fossor and T. pictus, and Liusus and Ht. crassus
synapomorphies supporting node 4. Nearly all of the (Fig. 1), and the monophyly and extreme morpho-
other named supraspecific groupings are also shown logical similarity of the two Liusus species, instead
to be monophyletic, and so in this group a phylogeny provide stronger arguments in favour of the status
based on diverse morphological characters, including quo.
characters not originally used to define these group- Although several taxonomic problems have been
ings, supports the current taxonomy. resolved in this paper, some challenges for future
The systematic position of Ht. wakefieldi has been research clearly remain. For example, in none of the
questioned for some time (Newton, 1973; Klimasze- primary trees from the simultaneous analysis was C.
wski et al., 1996; Leschen et al., 2003), and my analy- maxillosus recovered as a monophyletic group. Recent
sis corroborates earlier views that this species should authors have included North American specimens in
be removed from Hadrotes (Fig. 12). Although there is the subspecies C. m. villosus (Smetana & Davies,
apparent character support for placement within 2000; Smetana, 2004; present study), and although
three different subtribes (Philonthina, Staphylinina, they are generally distinct in vestiture patterns, I
and Xanthopygina), some characters consistent with know of no unambiguous autapomorphy or diagnostic
placement in Staphylinina include a membranous character for C. maxillosus, beyond the distinct
postcoxal process, distinctly bilobed ligula, mouthpart median lobe shape. Despite its characteristic appear-
chaetotaxy, and wing venation characters (completely ance (e.g. Fig. 1F), it is also reliably externally diag-
separate MP3, MP4, and CuA veins: Solodovnikov & nosed from other maxillosus group species only by
Newton, 2005). The presence of empodial setae and lacking diagnostic characters of other species. A
more than one anterolateral mental setae in Ht. further complication is that C. m. villosus and C.
wakefieldi is also inconsistent with recent concepts of galapagensis formed a clade in five of the 11 primary
Philonthina (Schillhammer, 1998; Smetana & Davies, trees of the simultaneous analysis, a relationship
2000). Two mesepimeral ridges (one marginal, one supported by the distinctively apically flared appen-
discal; Fig. 9I, arrows) seem to be present only in the dix of the internal sac (ch74-1; Fig. 4C, D). Despite
primitive Arrowinini (Solodovnikov & Newton, 2005) this novel character state, the polytomy at node 16
and in Staphylinini, and are widely distributed suggests the current taxonomy of C. maxillosus
within the latter. All other staphylinine tribes either should remain intact for now; C. maxillosus clearly
have a single marginal ridge, or none at all (Xantho- deserves more thorough taxonomic study than was
linini). This suggests that the single mesepimeral possible here. Overall, the external and genitalic
ridge (ch27-1; Fig. 9K, arrow) is an uncommon similarity of the New Zealand species (node 17) and
derived state within Staphylinini and in combination the polytomy in the maxillosus group (node 16) may
with other synapomorphies for node 3 it would seem indicate recent or rapid speciation in these groups
to stand out as particularly good phylogenetic evi- that is probably related to the colonization of mar-
dence for maintaining Ht. wakefieldi in the Creophi- ginal habitats. These morphological and phylogenetic
lus complex. Nevertheless, the contrast between this features suggest that species in these groups may be
evidence and particularly the mandibular and genital fertile ground for further speciation and biogeographi-
differences between Ht. wakefieldi and the rest of the cal studies.
Creophilus complex suggests that a more inclusive The present analysis integrates a diverse character
analysis and detailed taxonomic study of H wakefieldi set to investigate the systematics of this group, but it
is required to further test the placement of this did not consider larval characters, which are widely
species. known to be phylogenetically informative (Solodovni-
The marine littoral species are shown to not form kov, 2007; Meier & Lim, 2009), nor molecular data.
a monophyletic group. The placement of these taxa The general lack of material prevented inclusion of
both character systems, but larvae have been peritremes enlarged medially, with posterior edge
described for C. maxillosus (e.g. Dajoz & Caussanel, produced into point (Fig. 2E, ptm); mesoventral inter-
1969), Ht. crassus (Moore, 1964), Hd. fossor (Shibata, mesocoxal process truncate or broadly angulate, with
1965), and T. pictus (Böving & Craighead, 1931). mesocoxae widely separated (Fig. 2F, mip); female
Those for three other C. erythrocephalus group with gonostyle articulated subapically on broadly
species have been recently collected or discovered in rectangular or subcolumnar second gonocoxite
collections (pers. observ.). Discovering the larvae of (Fig. 5D, sty).
Liusus, Ht. wakefieldi, and at least one other species
of the C. maxillosus group would allow further testing Description: Habitus: Generally large (10–28 mm)
of the phylogenetic conclusions presented here. black or partly colourful rove beetles (e.g. Fig. 1A–J),
with (five spp., e.g. Fig. 1F–J) or without (seven spp.,
e.g. Fig. 1A–E) variegated whitish-grey or yellowish-
TAXONOMY
brown vestiture. Both sexes are strongly polymorphic
THE CREOPHILUS COMPLEX in size, males generally larger, with larger heads and
Adult diagnosis: Species of the Creophilus complex longer mandibles (both allometric). Head (Fig. 2A, B).
can be recognized by the combination of: disc of head Head trapezoidal to subrectangular; strongly trans-
and pronotum glabrous; ventroscapal macrosetae verse; shorter than pronotum; narrower than prono-
absent or reduced; superior marginal pronotal carina tum in females, slightly wider than pronotum in
separate from inferior marginal carina along entire males; mostly glabrous dorsally (disc). Frons and
length, effaced apically (Fig. 2E, il, sl, Fig. 9J, right vertex coarsely but sparsely punctured; or impunc-
and middle arrows); inferior marginal pronotal carina tate; punctures when present less than half diameter
continuous with anterior marginal carina (Fig. 9J, of macrosetal puncture. Antennal fossae separated
left and middle arrows); mesepimeron with single from anterior margin of head by sclerotized strip
marginal ridge (Fig. 9K, arrow); male sternite IX sub- (Fig. 2B, atf). Dorsum of neck sparsely punctate to
truncate to shallowly emarginate apically, produced impunctate, punctures when present as above. Con-
into elongate process basally (Fig. 3Q); median lobe of figuration of head ridges as in Figure 2A, B: nuchal
aedeagus with large apicolateral sclerites (Fig. 3, as); ridge generally incomplete dorsomedially, close to
internal sac without ventral basomedian sclerotized infraorbital ridge laterally; dorsal and ventral basal
strip (as in Fig. 3K, bms). ridges obliterated laterally by anastomosing setifer-
ous punctures; postmandibular ridge absent; gular
Remarks: The following taxa are here included in the sutures narrowly separate; basal gular impression
Creophilus complex: Creophilus (12 species), Hadrotes not demarcated by anterior transverse ridge; infraor-
crassus, Ht. wakefieldi, Hadropinus fossor, Liusus bital ridge present; ventral basal ridge on postgena
(two species), Thinopinus pictus. separated from postoccipital suture, terminating at
gular sutures. Antennae (e.g. Fig. 13F). Antennae
11-segmented; clavate to subincrassate; longer than
GENUS CREOPHILUS LEACH head; insertions exposed dorsally. Antennomeres 1–6
Creophilus Leach, 1819: 172. Type species: Staphyli- shining, not pubescent; 7–11 densely pubescent; 1–3
nus maxillosus Linnaeus, 1758, fixed by original des- elongate; 1 longer than 2 and 3 together; 2–3 sub-
ignation according to action of Commission in Opinion equal, dilated distally; 4 globular; 5–10 increasingly
546 (ICZN, 1959). Blackwelder (1952: 111) listed transverse (10 widest), somewhat dorsoventrally flat-
synonymic homonyms and variant spellings for tened; 11 narrower than preceding, dorsoventrally
Creophilus. compressed apically forming distinct ridge (except C.
rekohuensis), and bearing two pairs of prominent
Saprophilus Streubel 1839: 136; Blackwelder 1952: setae. Mouthparts. Labrum (Fig. 2B) short and trans-
344 (synonym of Staphylinus). Type species: Staphyli- verse; broadly incised mesially; partly sclerotized, and
nus maxillosus Linnaeus, 1758, fixed by monotypy. with numerous macrosetae. Mandibles falciform; each
with distinct mandibular blades (Fig. 2D, mb); with
Adult diagnosis: Within the Creophilus-complex, the dorsal basolateral ridge bordered by setiferous groove
following characters are exclusive to Creophilus: removed from edge (Fig. 2D, dbl). Right mandible
antennae clavate with antennomeres 7–10 much with two dorsal teeth (Fig. 2D, T1, T2) and one
wider than long (e.g. Fig. 13F) and only 7–11 densely ventral tooth (Fig. 2D, T3). Left mandible with two
pubescent; left mandible with distinct ventral baso- teeth in same plane (Fig. 2D); with distinct ventral
lateral ridge (Fig. 2C, vbr); protibiae with two spines basolateral ridge bordered laterally (especially in
on inner face (Fig. 9E, arrows); sternacostal ridge large specimens) by distinct groove to receive right
broadly arcuate medially (Fig. 2E, sr); mesothoracic mandible (Fig. 2C, vbr). Maxillae as in Figure 2H.
756 D. J. CLARKE
Labium as in Figure 2I. Mentum short, transverse; sinuous laterally, straight medially (Fig. 2L, btf). All
with two pairs of anterolateral macrosetae. Ligula segments with distinct basal transverse fold dorsally
minutely incised apically. Thorax. Pronotum trans- and ventrally, extending onto inner parasclerites
verse; strongly transversely convex; glabrous (on (Fig. 2L, btf). Tergites and sternites arcuate at
disc), shining; without definite microsculpture on disc; middle; with shallow transverse lateral impressions.
with anterolateral declivities, sides, and base sparsely Tergite I setose laterally; without prototergal glands,
to rather densely setose. Basolateral pronotal but sometimes with shallow acetabula laterally.
margins faintly to distinctly emarginated. Pronotal Tergite II as in Figure 2J; with short internal ante-
hypomeron visible in lateral view; glabrous. Pronotal rolateral flanges (af); with paired muricate fossae (f).
postcoxal process large and triangular; hyaline; fim- Second abdominal spiracle large; situated in mem-
briate anteriorly. Superior pronotal marginal line brane outside of tergite. Apical margin of tergite VII
separate from inferior line, never joining it anteriorly; with distinct white palisade fringe in all macropter-
deflexed beneath anterior pronotal angles, thus ous species (e.g. Fig. 1A; reduced or absent in flight-
pronotum anteriorly visible in ventral view (Fig. 2E). less species). Paired abdominal defensive gland
Inferior pronotal marginal line continuous with ante- reservoirs present (for detail see Dajoz & Caussanel,
rior pronotal margin (Fig. 2E). Prosternum (Fig. 2E) 1969: fig. 10A). Male. Head as wide as or slightly
fused with hypomeron (pronotosternal suture absent). wider than maximum pronotal width. Sternite VIII
Basisternum broadly rounded; sparsely setose; acari- deeply subtriangularly emarginate (e.g. Fig. 23K).
nate; without distinct intercoxal depressions. Sterna- Apices of tergite X and sternite IX subtruncate to
costal ridge broadly arcuate medially. Furcasternum shallowly emarginate, and sternite IX with elongate
glabrous; almost vertical; with distinct fovea; poste- basal portion (Fig. 3Q). Aedeagus (e.g. Fig. 3A–C).
riorly with broad groove connecting posterolateral Aedeagus bilaterally symmetrical; with paramere
fossae (Fig. 2E, plf). Mesothoracic peritremes very facing left in repose (in abdomen). Apex of median
large and acute; approximate mesially; with rein- lobe with paired apicolateral sclerites (as); deeply
forced poriferous membrane behind (Fig. 2E, ptm). incised laterally, proximal to lateral recurved teeth
Scutellum densely punctured; setose. Elytra. Elytra (lit; except in C. albertisi). Basal bulb with large
together subrectangular; longer than pronotum (when dorsal sclerite, and paired ventral sclerites separated
measured from humerus to apex); with epipleura medially by membrane. Paramere spatulate; nar-
faintly impressed anteriorly; with subapical surface rower than median lobe; shaft widest apically or
depressed laterally. Apices obliquely truncate; preapically; not extending beyond apex of median
forming obtuse angle with suture. Disc and epipleu- lobe; emarginate apically (except C. variegatus); with
ron evenly setose, but without micropubescence; several apicolateral setae; dorsoventrally movable at
without microsculpture (but frequently rugosely base (i.e. not fused to median lobe; except C. alber-
sculptured). Humeri sparsely punctate and setose to tisi). Internal sac with structures as labelled in
impunctate and at least partly glabrous; often with Figure 3A–D; bilaterally symmetrical, everting sym-
distinctive shining humeral callus (e.g. Fig. 1A). metrically along axis of median lobe (Fig. 3C, D).
Wings. Wings well developed and functional (except C. Membranous parts of internal sac, especially distally
rekohuensis and C. huttoni); with MP3, MP4, and around copulatory piece, armed with minute tooth- or
CuA veins completely separate (Fig. 33N). Mesoven- hair-like scales (Fig. 4, sh). Female. Head as wide as
trite acarinate; with short broadly transverse inter- or narrower than maximum width of pronotum.
mesocoxal process (Fig. 2F, mip). Metaventrite Sternite VIII entire. Apices of tergite X (except C.
broadly convex; with marginal carinae of mesocoxal variegatus) subtruncate to shallowly emarginate.
acetabula obsolete posteromedially (Fig. 2F, mac); Gonocoxites I and II (Fig. 5D, gc) separated by mem-
with deep premetacoxal groove (Fig. 2F, pmg). Legs. brane. Distal gonocoxites broadly rectangular or sub-
Tibiae multispinose, with numerous longer bristles columnar; with gonostyle (sty) articulated subapically
interspersed (e.g. Figs 2K, 9E). Tarsi pentamerous; (i.e. articulation point not visible in ventral view).
with bisetose empodia. Prothoracic legs robust. Internal female genitalia (e.g. Fig. 5A–C). Vaginal
Protibiae (Fig. 2K) moderately expanded distally; cavity with structures as labelled in Figure 5A–C.
spinose only posteriorly; with two prominent spines Spermatheca unsclerotized, tubular to only very
on ventral face (opposing profemur when leg slightly expanded distally. Chaetotaxy (Fig. 6). All
adducted; Fig. 9E). First four protarsomeres trans- named macrosetae, when present, represented by
versely bilobed in both sexes; clothed ventrally with single macrosetae, never clusters (excluding procoxal
tenent setae. Mesocoxae widely separated. Abdomen. and posterior elytral ‘patches’: pp, pt), and are illus-
Abdomen densely setose, with setae moderately trated for a hypothetical member of the Creophilus
appressed, and sparser or absent anteriorly and complex in Figure 6. (Head) Anteromarginal (am)
around spiracles. Basal tergal margins slightly anterior to antennal insertion; temporal and occipital
at posterior margin of head (tm, oc); ventroscapal (vs) Arrival and departure times for adults at carrion
same size as and indistinguishable from other setae, are highly variable, depending on environmental
or else absent. (Prothorax) Pronotum at most with conditions affecting decomposition rates (Archer,
only antero- and basolateral (an, bp); without anterior 2004).
pronotal group (ap). Procoxa with mesal patch (pp);
protrochanter with at least one (tr); profemur with
none. (Pterothorax) Elytra with at least one baso- Nomenclature and taxonomy: Samouelle (1819) pub-
humeral (bh); elytral discal series comprising two to lished the new generic name Creophilus, for which he
eight (ds); posterior elytral patch with at least three designated Staphylinus maxillosus Linnaeus, 1758 as
(pt); mesoventrite with broadly arcuate indistinct type species. This designation was later formally
row (mr); mesocoxae ventrally, and hind legs, with established in Opinion 546 (ICZN, 1959); Leach can
none. Metaventrite with one posteromedially (pm). be inferred as author of the genus by similar logic to
(Abdomen) Abdominal tergals placed at apical that of Opinion 1722 (ICZN, 1993). Since 1819, Creo-
margins on tergites III–VI, nearer middle on tergites philus has been considered either as a valid genus or
VII and VIII (ol, il, mm); parasclerites with none; a synonym or subgenus of Staphylinus (see Smetana
outer laterals (ol) present on all tergites. & Davies, 2000). Nordmann (1837) described C. badi-
ipennis, C. erithacus, and C. chloris, but these were
Distribution: Worldwide, but apparently absent from removed to the xanthopygine genus Torobus Herman
sub-Saharan Africa. Several species have become (Herman, 2001a). Staphylinus luctuosus Blanchard,
established outside their native ranges. 1842 was transferred to Creophilus by Bernhauer &
Schubert (1914), but Newton (1996) transferred it to
Biology and ecology: The biology and life history of Xanthopygus Kraatz after examination (and designa-
C. maxillosus are well known (see references in tion) of the lectotype.
Herman, 2001b). Comparable mouthpart morphology The nomenclature of C. maxillosus-group species is
and specimen data suggest that all species (except marked by proliferation of specific, infraspecific, and
possibly C. huttoni) are primarily predators of infrasubspecific synonyms, reflecting variability
dipteran larvae at carrion, although they may occa- within some species. Recent works (e.g. Smetana &
sionally be found in other decaying substrates, or Davies, 2000; Smetana, 2004) treat C. m. maxillosus
take other prey. Facultative cannibalism has also (Linnaeus) and C. m. villosus (Gravenhorst) as sub-
been observed (Fichter, 1948). They are not nor- species, and this classification is maintained here.
mally necrophagous and prefer carrion in the early Types of 12 of the 20 subjective C. maxillosus syn-
stages of decomposition (Fichter, 1949a) when onyms were studied, and most synonyms have been
blowfly larvae are most plentiful. A preference for allocated to the appropriate subspecies. Several
blowfly larvae has been demonstrated for C. m. vil- names were also determined to be unavailable. A
losus adults and larvae in feeding experiments Galapagos Islands species, frequently referred to as
(Abbott, 1934; Fichter, 1948; Greene, 1996), and ori- C. m. villosus, is here described as the new species C.
entation to the carrion resource is achieved using galapagensis.
long-range olfactory cues (Abbott, 1934, 1936, 1938). Nomenclatural issues in the C. erythrocephalus-
One consequence of the carrion microhabitat asso- group are fewer, involving confusion between C. eryth-
ciation is that adults can be collected en masse rocephalus, C. lanio, and C. oculatus, or disagreement
using carrion-baited pitfall traps. Single squid- in their rank. However, both Fauvel (1875) and Steel
baited traps (Newton & Peck, 1975) left for several (1949) clearly distinguished them at the specific level.
weeks can collect hundreds of specimens. In con- For the New Zealand region, Herman (2001b) listed
trast, larvae and pupae are evidently never only one valid species (C. oculatus), but Emberson
collected using this method, but can be collected (1998) suggested that C. oculatus tends to be replaced
in the vicinity of carrion (pers. observ.). This by another species in the south of South Island. This
suggests that larvae hatch from eggs laid directly on distinctive beach-inhabiting species (C. huttoni),
or near the carrion following arrival of adults from known also to Newton (1985) and Hammond (2000),
elsewhere. The likely requirement that the life cycle was synonymized by Gourlay (1950) without
be completed in situ only at carrion or similar sub- comment. Moreover, Emberson (1998) also reported a
strates might explain the absence of larvae from species from the New Zealand Chatham Islands indi-
baited traps. Pre-adult development takes up to 35 cating that it was ‘very similar or identical to the one
days in C. maxillosus (details in Kramer, 1955), so [C. huttoni] found in the southern South I. (SL) and
newly emerged adults will probably disperse due to Stewart I.’ Creophilus huttoni is here reinstated,
local conditions unsuitable for feeding and develop- and C. rekohuensis is described from the Chatham
ment (carrion dried out, low maggot abundance). Islands.
758 D. J. CLARKE
KEY TO ADULTS OF CREOPHILUS SPECIES

(1) Head black; first three visible abdominal tergites (i.e. III–V) without secondary basal transverse ridge; body
vestiture variably black mixed with yellowish- or whitish-grey (maxillosus-group, e.g. Fig. 1F–J) ............... 2
(1⬘) Head orange-red or yellowish-brown at least ventrally; first three visible abdominal tergites (i.e. III–V) with
secondary basal transverse ridge (Fig. 2L, sbr); body vestiture uniformly black or brown (erythrocephalus-group,
e.g. Fig. 1A–E) .............................................................................................................................7
(2) Antennomere 11 convex apically; anterior lengths of antennomeres 9 and 10 distinctly less than posterior
lengths (Figs 16B, 23B) ................................................................................................................. 3
(2⬘) Antennomere 11 concave apically; anterior lengths of antennomeres 9 and 10 subequal to posterior lengths
(Figs 13F, 18B, 21C) ..................................................................................................................... 4
(3) Fifth visible abdominal tergite (i.e. VII) with extensive whitish setae, especially apicolaterally; elytral cuticle
bicoloured, disc reddish-brown, sides and humeri yellowish; elytra with variegated golden, brown, and
whitish setae; basiantennal macroseta absent; basolateral pronotal macroseta present; South America
...................................................................................................................(5) variegatus (Fig. 1J)
(3⬘) Fifth visible abdominal tergite (i.e. VII) with mostly or entirely black setae; elytral cuticle uniformly black or
brownish-black; elytra with only silvery-whitish and black setae; basiantennal macroseta present; basolateral
pronotal macroseta absent; Madagascar, Réunion, Mauritius, Mayotte ....................... (2) incanus (Fig. 1G)
(4) Elytral setae uniformly black; abdominal surface between punctures appearing dull, with imbricate microscu-
lpture (Fig. 19A); abdominal tergites III–VII each with only two macrosetae, one at each side; Galápagos Islands
..................................................................................................(3) galapagensis sp. nov. (Fig. 1H)
(4⬘) Elytral setae mostly whitish, silvery-grey, or yellowish, and black; abdominal surface between punctures
appearing shiny, with transversely strigulate microsculpture (Fig. 19B); most abdominal tergites with at least
six macrosetae .............................................................................................................................5
(5) Fringe of setae at apices of first four visible tergites (i.e. III–VI) with mostly or entirely whitish-grey or yellowish
setae; elytral vestiture mostly whitish-grey or yellowish; abdomen dorsally with mostly black setae; eyes very
large, occupying most of sides of head (HL1/HL2 usually greater than 4.0, range 1.92–12.0; e.g. Fig. 21A);
widespread in Oriental Region ........................................................................ (4) flavipennis (Fig. 1I)
(5⬘) Fringe of setae at apices of first four visible tergites (i.e. III–VI) with paired groups of whitish-grey or yellowish
setae near middle; elytra usually with distinct angulate fascia of whitish- or silvery-grey setae (Fig. 13B);
abdomen dorsally with extensive whitish-grey or yellowish vestiture at least on tergites IV–V (Fig. 1F); eyes
much smaller, with well-developed temples (HL1/HL2 usually less than 3.0, range 1.21–3.60); Holarctic,
Neotropical ................................................................................................................................. 6
(6) Vestiture of hind margins of head, anterior pronotal angles, and pterothorax (Fig. 14A) mostly black; abdomen
with extensive whitish-grey vestiture on most or all tergites; Palearctic, north-west Nearctic, probably adventive
in east Nearctic, South America ....................................................(1a) maxillosus maxillosus (Fig. 1F)
(6⬘) Vestiture of hind margins of head, anterior pronotal angles, and pterothorax (Fig. 14B) whitish-grey; abdomen
with extensive whitish-grey vestiture on tergites and parasclerites IV–V only; USA, Canada, Mexico, Central
America, West Indies, Hawaii (probably adventive) ...........................................(1b) maxillosus villosus
(7) Elytra with metallic blue-purple or blue-green luster ..........................................................................8
(7⬘) Elytra uniformly black or brown .....................................................................................................9
(8) Head with circular black spot at centre of vertex (Fig. 7A), without posterolateral tooth; antennomeres 7–11
greyish-black or black; abdominal segments VIII and IX uniformly black; Australia, widespread across Pacific
.......................................................................................................... (6) erythrocephalus (Fig. 1A)
(8⬘) Head with triangular black spot continuous with frontal margin (Fig. 7E), with posterolateral tooth (Fig. 31A,
B, vlc); antennal segments 7–11 whitish-orange; abdominal segments VIII and IX bright orange; New Guinea
......................................................................................................................(9) albertisi (Fig. 1E)
(9) Head orange, with black spot at centre of vertex (Fig. 7B, D); protrochanter with one macroseta, mesotrochanter
without macrosetae; Australia and New Guinea ...............................................................................10
(9⬘) Head with vertex (between eyes) entirely black or blackish-brown, or mottled yellowish-brown and brownish-
black (Figs 1B–D, 7C); protrochanter with two macrosetae, mesotrochanter with one macroseta; New Zealand
subregion .................................................................................................................................. 11
(10) Medial black spot small, circular, sharply defined (Fig. 7D); abdominal segment IX black; pronotum with smooth
basolateral impressions (Fig. 27E); right mandible with two distinct teeth (Fig. 27A); New Guinea ...............
.................................................................................................................................. (7) imitator
(10⬘) Medial black spot large, variable in shape, oblong-subquadrate or cordiform, edges diffuse (e.g. Fig. 7B);
abdominal segment IX orange-brown; pronotum without basolateral impressions (Fig. 29F); right mandible with
three distinct teeth (Fig. 29A); Australia ................................................................................(8) lanio
(11) Antennomere 11 without distinct apical ridge, asymmetrically produced to anterior point (Fig. 36B, arrow);
apical protarsomere shorter than preceding two together (Fig. 33K); protibiae with between zero to (rarely) three
short spines (Fig. 33L); body uniformly brownish-black dorsally, head yellowish or orange-red only ventrally;
Chatham Islands .......................................................................... (12) rekohuensis sp. nov. (Fig. 1D)
(11⬘) Antennomere 11 with distinct apical ridge (e.g. Fig. 32B, arrow); apical protarsomere longer than preceding two
together (Fig. 33J); protibiae with numerous (always > 3) long thick spines (Fig. 33M); body variably black or
reddish-brown to brownish-black, with or without reddish or yellowish spots behind eyes ...................... 12
(12) Fifth visible abdominal tergite (i.e. VII) with distinct white palisade fringe (Fig. 1C); mesoventral intermesocoxal
process obtuse medially (Fig. 33G); macropterous (Fig. 33N); body black, head with distinct orange-red spots
posterior to eyes (Fig. 7C); widespread ............................................................... (10) oculatus (Fig. 1C)
(12⬘) Fifth visible abdominal tergite (i.e. VII) without or with only vestigial palisade fringe (Fig. 1B); mesoventral
intermesocoxal process angulate medially (Fig. 33H); brachypterous (Fig. 33O); body variably reddish-brown to
brownish-black (rarely mostly black); head usually with yellowish (rarely red or without) spots behind eyes;
South Island, Stewart Island, marine littoral ........................................................(11) huttoni (Fig. 1B)
Figure 13. Creophilus maxillosus, details of morphology (setae of mandibles, antenna, and pronotum omitted). C. m.
maxillosus (A–G); C. m. villosus (H). A, mandibles, dorsal (bases omitted); B, right elytron (only whitish setae shown) C,
paramere, dorsal; D, aedeagus with details of inverted internal sac, right lateral (parameral setae omitted); E, pronotum
(small male); F, right antenna, dorsal (only apical setae shown); G, female internal genitalia, ventral H, ventral sclerite
of internal sac, ventral (lower) and apical (upper). Abbreviations: T1, T2, T3, mandibular teeth. Additional abbreviations
as in Figure 3 (male) and Figure 5 (female). Scale bars: H = 0.1 mm; rest = 1 mm.
CREOPHILUS MAXILLOSUS SPECIES-GROUP Diagnosis: Head black or brownish-black; antenno-

This species group was originally proposed by Newton mere 2 subequal in length to 3 or shorter; antennomere
(1985) for the Holarctic and Oriental species. With 11 with apical pairs of setae situated on either side of
recognition of three new synonyms and description of antennal axis (e.g. Fig. 13F); right mandible with T3
one new species, five species are included here. directly ventral to T2 and mandibular blade, mostly (in
760 D. J. CLARKE
C. incanus entirely) hidden from dorsal view beneath of C. maxillosus s.l.); Herman, 2001b: 3323
mandibular blade (Fig. 13A); anterior pronotal angles (synonym of C. maxillosus s.l.; preoccupied by Sta-
produced laterally (Figs 2G, 13E); abdominal tergites phylinus bicinctus Rossi, 1792); Navarrete-Heredia
III–V without secondary transverse basal ridge; apical et al., 2002: 322 (synonym of C. maxillosus s.l.);
fringe of tergites III–VI with at least paired groups of Smetana, 2004: 671 (synonym of C. m. maxillosus);
white or golden setae medially, rest of body with primary junior homonym of Staphylinus bicinctus
variably black, brown, whitish-grey, or golden-brown Rossi, 1792.
vestiture (Fig. 1F–J); median lobe of aedeagus com- Staphylinus orientalis Motschulsky, 1858b: 67.
pletely divided ventrally by membranous strip Type locality: ‘Iles Kouriles’; Ganglbauer, 1895: 415
(Fig. 3C, arrow). Large males with: apical portion of (synonym of C. maxillosus s.l.); Herman, 2001b: 3324
mandibles exceeding 0.9 times length of basal portion (synonym of C. maxillosus s.l.); Smetana, 2004: 672
(ML1/ML2 range of all males = 0.60–1.09); without (synonym of C. m. maxillosus).
ventrolateral carina of head; pronotum widest at ante- Creophilus fulvago Motschulsky, 1860: 120. Type
rior angles; base of profemur with spine-like apophysis locality: ‘Mongolie et Chine boréale’; Ganglbauer,
(Fig. 9D, arrow); superior pronotal marginal line 1895: 415 (synonym of C. maxillosus s.l.); Herman,
abruptly deflected beneath disc of pronotum at point of 2001b: 3324 (synonym of C. maxillosus s.l.); Smetana,
basolateral denticle (Fig. 2G, bpd). 2004: 672 (synonym of C. m. maxillosus).
Creophilus maxillosus var. pulchellus Meier, 1899:
99. Type locality: ‘Hamburg’; Blackwelder, 1943: 448
1. CREOPHILUS MAXILLOSUS (LINNAEUS)
(synonym of C. maxillosus s.l.); Herman, 2001b: 3324
(FIGS 1F, 2E–G, 3C, D, I, P, 4B, C, 5A, 9C–F, L, (synonym of C. maxillosus s.l.); Smetana, 2004: 672
M, 10B, 13, 14, 15,19B, 24) (synonym of C. m. maxillosus).
Staphylinus maxillosus Linnaeus, 1758: 421. Type
locality: ‘Europa’; Herman, 2001b: 3317 (as C. maxil- Diagnosis: With characters of the maxillosus-group;
losus); Navarrete-Heredia et al., 2002: 322 (as C. max- integument, including elytra, uniformly black; anten-
illosus). Type specimens: see C. maxillosus nae moderately clavate; apex of antennomere 11
maxillosus. emarginate (Fig. 13F); elytra with white vestiture
Staphylinus anonymus Sulzer, 1761: 17, fig. 49. arranged into well-demarcated (but variable) central
Type locality: ‘Europa’; Erichson, 1839: 348 (synonym transverse fascia (Figs 1F, 13B); tergal chaetotaxic
of S. maxillosus s.l.); Herman, 2001b: 3321 (synonym formula 6-6-6-2(4)-4(6)-8(10+).
of C. maxillosus s.l.); Smetana, 2004: 671 (synonym of
C. m. maxillosus). Description: To previous detailed descriptions (Black-
Staphylinus tertius Schaeffer, 1766: plate 20, fig. 1, welder, 1936; Dajoz & Caussanel, 1969) I add mea-
1. Type locality: ‘Ratisbon’ (not seen); Erichson, 1839: surements and body ratios (listed under each
348 (synonym of S. maxillosus s.l.); Herman, 2001b: subspecies), chaetotaxic characters, and full descrip-
3321 (synonym of C. maxillosus s.l.; nomen nudum, tion of male and female genital characters. Head.
Article 11.4). Head slightly wider anteriorly in large males, sub-
Staphylinus balteatus DeGeer, 1774: 18. Type local- trapezoidal and wider posteriorly to suborbicular in
ity not cited; Fabricius, 1781: 334 (synonym of S. females and smaller males; basal margins densely
maxillosus s.l.); Herman, 2001b: 3321 (synonym of C. setose; dorsal punctation moderately dense; eyes
maxillosus s.l.); Smetana, 2004: 671 (synonym of C. large, dorsolateral, lateral margins of head visible in
m. maxillosus). dorsal view (not obscured by eye), HL1/HL2 greater in
Staphylinus fasciatus Füessly, 1775: 21. Type local- females than males; antennae as in Figure 13F, mod-
ity: Switzerland; Gravenhorst, 1802: 3 (synonym of S. erately clavate; antennomeres 1–6 brownish-black,
maxillosus s.l.); Herman, 2001b: 3322 (synonym of C. 7–11 greyish-black; apex of antennomere 11 slightly
maxillosus s.l.); Smetana, 2004: 672 (synonym of C. to moderately emarginate medially; each pair of
m. maxillosus). apical setae widely separated, one on each side of
Staphylinus nebulosus Geoffroy, 1785: 165. Type apex; mandibles as in Figure 13A, T1 and T3 sub-
locality: Paris; Olivier, 1795: (42): 10 (synonym of S. equal in length, in large males T1 much larger than
maxillosus s.l.); Herman, 2001b: 3322 (synonym of C. T2 and T3. Thorax and abdomen. Pronotum (Figs 2G,
maxillosus s.l.); Smetana, 2004: 672 (synonym of C. 13E) moderately to distinctly narrowed posteriorly;
m. maxillosus). basolateral margins very shallowly emarginate, hind
Staphylinus bicinctus Mannerheim, 1843: 229. angles indistinct; with dense and long peripheral
Type locality: ‘insula Sitkha’; Solsky, 1870: 261 setae and coarse long black or whitish-grey vestiture
(synonym of C. arcticus); Horn, 1879: 200 (synonym on anterolateral declivities; scutellum with indistinct
of C. villosus); Ganglbauer, 1895: 415 (synonym anterior band of whitish-grey vestiture; elytra uni-
formly black, densely setose, with characteristic well- Distribution: Widespread throughout the northern
delimited fascia of whitish-grey setae at middle; hemisphere; apparently absent from austral regions
wings fully developed, clear yellowish-brown to black, (except as adventive), South East Asia, and the East
without black spot in medial field between MP3 Indies. See under each subspecies for details.
and MP4 veins; ventral thoracic vestiture entirely
whitish-grey or black (Fig. 14); anterior and middle Biology and ecology: Creophilus maxillosus is prob-
legs with black vestiture, hind coxae, trochanters, and ably the best-studied rove beetle species. Herman
femora with white vestiture; abdominal vestiture (2001b) lists many papers on its biology, morphology,
arranged into characteristic pattern of whitish-grey and life history. The following papers provide details
or golden brown maculation dorsally, and mostly or of: general biology and life history (Abbott, 1938;
entirely white on sternites and parasclerites IV–VI, Fichter, 1949b; Kramer, 1955; Greene, 1996);
sternite III, and sternites VII–IX; extensive whitish- morphology and anatomy (Talbot, 1928; Blackwelder,
grey vestiture interrupted dorsally by a large medial 1936; Dajoz & Caussanel, 1969); diet (Fichter, 1948);
and two smaller anterolateral black patches, and ven- chemical defence (Jefson et al., 1983); olfaction
trally by more or less circular black patches laterally; (Abbott, 1936); and immature morphology (Voris,
abdominal tergite VII with well-developed palisade 1939; Paulian, 1941). Kirkpatrick & Olson (2002) note
fringe. Male genitalia. Aedeagus as in Figures 3C, increases in predatory activity at night. Moon & Kajii
13D; paired apicolateral sclerites articulated to (1997) observed fully grown larvae feeding on fly
internal edge of median orifice. Paramere as in maggots beneath rabbit carcasses, where they are
Figure 13C. Internal sac inverted as in Figure 13D, presumably able to evade the cannibalistic tendencies
everted as in Figure 3C, D; with short ventral process of adults (Abbott, 1937, 1938) by burrowing into the
(vp), ventromedian spiculose strip slightly longer substrate (as observed by Fichter, 1948).
than the box-like ventral sclerite (Figs 3I, 13H, vs). Dynamics of arrival and departure at carrion have
Female internal genitalia. Internal genitalia as in been recorded frequently in the forensic entomology
Figures 5A, 13G; vaginal plate produced into two literature (e.g. Early & Goff, 1986; Goff, 1991; Ander-
broad lobes posteriorly, and with median sclerotized son & VanLaerhoven, 1996; Moon & Kajii, 1997; Rich-
strip (ss); vaginal fold (vf) completely membranous; ards & Goff, 1997; Sánchez Piñero, 1997; Avila & Goff,
spermatheca (sp) elongate sacculate, with distinct 1998). Summarizing these papers, adults arrive early
duct connecting to bursa copulatrix (bcx). Chaetotaxy. at a carcass during the ‘bloat’ and ‘decay’ stages
Elytral discal series with 4–8 macrosetae, frequently (Payne, 1965; length of stages varies depending on
with additional macroseta medially disposed from environmental conditions), remaining there up to
series; tergal chaetotaxic formula = 6-6-6-2(4)-4(6)- 13 days post-mortem. Creophilus maxillosus prefers
8(10+), outer laterals present or absent and inner open disturbed habitats over dense forest, and is
laterals absent on tergite VI, inner laterals present or typically found in synanthropic situations. Habitat:
absent on tergite VII; tergite VIII often with incon- seashore, grassland, prairie, mixed forest, savannah,
sistent additional macrosetae. woodland, swamp, alpine meadow, etc. Altitude: sea
level to 3640 m. Phenology: throughout the year,
Variation: Coloration and the patterning of body ves- tending to be most common from late spring through
titure are the only notable geographically variable autumn in northern USA.
characters, and with the exception of North American
and Palearctic forms (discussed under each subspe- Remarks: Measurement and ratio data for this species
cies) appears to be continuous. These forms exhibit are provided for each subspecies, below. I could not
more or less identical male and female genitalia, determine details and/or depositions for type material
with only subtle differences in the copulatory piece/ of the following synonyms: Staphylinus anonymus
appendix that may be significant for reproductive Sulzer, Staphylinus tertius Schaeffer, Staphylinus
isolation but which require more detailed study. Much balteatus De Geer, Staphylinus fasciatus Füessly, Sta-
of the synonymic diversity is attributable to varia- phylinus nebulosus Geoffroy, Staphylinus bicinctus
tions in vestiture coloration and to allometric charac- Mannerheim (syntypes in ZMHB?), and C. maxillosus
ters of the head and thorax. var. pulchellus Meier (holotype by monotypy). Syn-
types (?) of S. orientalis and C. fulvago are presum-
Comparison: Creophilus maxillosus is distinctive in ably in ZMUM but according to A. Gussakov (ZMUM)
habitus (Fig. 1F) and most similar to C. incanus. these specimens are in poor condition and their iden-
It is reliably distinguished from this species by the tity cannot be confirmed. I list these nine names here
concave apex of the apical antennomeres, and from all because I could not confirm their status as synonyms
species by the definite whitish-grey elytral fascia, and of the nominate subspecies, which for most is never-
dorsal pattern of body vestiture. theless suggested by their type localities and syn-
762 D. J. CLARKE
onymic history. All other synonyms and type material Creophilus maxillosus var. imbecillus Sharp, 1874:
are listed under the subspecies to which they apply. 28. Type locality: ‘Japan’; Ganglbauer, 1895: 415
Previous authors (e.g. Horn, 1879; Hatch, 1957) have (synonym of C. maxillosus s.l.); Herman, 2001b: 3324
treated C. bicinctus as a synonym of C. m. villosus (synonym of C. maxillosus s.l.); Smetana, 2004: 672
or C. arcticus (see Hatch, 1938: 148). Motschulsky (synonym of C. m. maxillosus); (unavailable: infra-
(1858b, 1860) notes the similarity of S. orientalis to C. subspecific name, Article 45.6.4).
m. maxillosus, noting that it differs only in (trivial) Creophilus maxillosus ssp. canariensis Bernhauer,
dimensions of the head and thorax. According to the 1908: 334. Type locality: ‘kanarische Inseln’; Black-
original descriptions, C. fulvago Motschulsky and C. welder, 1943: 448 (synonym of C. maxillosus s.l.);
m. var. pulchellus Meir may represent forms similar Herman, 2001b: 3324 (synonym of C. maxillosus s.l.);
to the C. ciliaris holotype, characterized by extensive Smetana, 2004: 672 (synonym of C. m. maxillosus).
golden or reddish-brown vestiture over most of the Creophilus sikkimensis Wendeler, 1927: 8. Type
body. Most authors prior to 1938 treated C. villosus locality: ‘Sikkim’; Herman, 2001b: 3325; Schillham-
(Gravenhorst, 1802) as either a separate species or a mer, 2004: 324 (synonym of C. maxillosus s.l.);
variety of C. maxillosus (Herman, 2001b). Hatch Smetana, 2004: 672 (synonym of C. m. maxillosus).
(1938) was the first to treat C. villosus as a subspecies
of C. maxillosus, although Blackwelder (1943) later Type material: Staphylinus maxillosus Linnaeus. Syn-
synonymized it with C. maxillosus, which was fol- types (?) presumably in UZIU (Blackwelder, 1943) (not
lowed by Herman (2001b). The subspecific division of seen).
C. maxillosus has been readopted in some recent Creophilus ciliaris Stephens. Holotype (by monotypy).
works (Smetana & Davies, 2000; Smetana, 2004), 씹 (BMNH, not seen). According to Mr R. Booth (BMNH),
including this one, because although superficial, the holotype is ‘pin-staged with a label “ciliaris” in brown
the differences between them are more or less dis- ink. The handwriting is W.E. Leach’s but Leach had left
crete, and may be supported by some internal sac the Museum before Stephens published his description,
differences that require further study. so cannot have labeled another individual after the
description was published.’ (in litt., 7 June 2006). The
1A. CREOPHILUS MAXILLOSUS MAXILLOSUS
specimen has been restaged on a more secure mount,
transferred to a drawer at the end of the Stephens
(LINNAEUS)
collection, and labelled with a red-bordered, circular
(FIGS 1F, 2E–G, 3C, D, I, P, 4B, 13A–G, BMNH Holotype label, and ‘HOLOTYPE Creophilus
14A, 15, 24) ciliaris Stephens, 1832: 202 det R.G. Booth 2006’. He
Staphylinus maxillosus Linnaeus, 1758: 421. Type notes that ‘apart from the golden pubescence, instead
locality: ‘Europa’; Leach, 1819: 172 (type species of of the more usual dark brown/black pubescence on the
Creophilus); Herman, 2001b: 3317 (as C. maxillosus). apices of the elytra and tergites, it looks like any other
Creophilus ciliaris Stephens, 1832: 202. Type local- C. maxillosus’.
ity: ‘Edinburgh’; Kraatz, 1857: 529 (synonym of C. Staphylinus cinerarius Erichson. Lectotype (here
maxillosus s.l.); Herman, 2001b: 3323 (synonym of C. designated). 씹, ‘[orange] Type/ 5878/ [green] ciner-
maxillosus s.l.); Smetana, 2004: 672 (synonym of C. arius| Er.| Aegypt xiii. 13. Ehbg/ [white] Hist.-Coll.
m. maxillosus). (Coleoptera)| Nr. 5878| Staphylinus cinerarius|
Staphylinus cinerarius Erichson, 1839: 350. Type Erichs.| Aegypt., Ehrenberg| Zool. Mus. Berlin/ [red]
locality: ‘Aegypto’; Blackwelder, 1943: 448 (synonym SYNTYPUS| Staphylinus| cinerarius Erichson,
of C. maxillosus s.l.); Herman, 2001b: 3323 (synonym 1839| labeled by MNHUB 2004/ FMNH-INS
of C. maxillosus s.l.); Smetana, 2004: 672 (synonym of 0000 016 786/ [red] LECTOTYPE| Staphylinus| cin-
C. m. maxillosus). erarius Erichson, 1839| designated by| D. J. Clarke
Creophilus maxillosus var. subfasciatus Sharp, 2008’ (in ZMHB). Paralectotypes (2). 씹, ‘[orange] Type/
1874: 28. Type locality: ‘Japan’; Ganglbauer, 1895: [white] Hist.-Coll. (Coleoptera)| Nr. 5878| Staphyli-
415 (synonym of C. maxillosus s.l.); Herman, 2001b: nus cinerarius| Erichs.| Aegypt., Ehrenberg| Zool.
3324 (synonym of C. maxillosus s.l.); Smetana, 2004: Mus. Berlin/ [red] SYNTYPUS| Staphylinus| ciner-
672 (synonym of C. m. maxillosus); (unavailable: arius Erichson, 1839| labeled by MNHUB 2004/
infrasubspecific name, Article 45.6.4). FMNH-INS 0000 016 787/ [yellow] PARALECTO-
Creophilus maxillosus var. medialis Sharp, 1874: TYPE| Staphylinus| cinerarius Erichson, 1839| des-
28. Type locality: ‘Japan’; Ganglbauer, 1895: 415 ignated by| D. J. Clarke 2008’ (in ZMHB); 씸, same
(synonym of C. maxillosus s.l.); Herman, 2001b: 3324 label data and ‘FMNH-INS 0000 016 788/ [yellow]
(synonym of C. maxillosus s.l.); Smetana, 2004: 672 PARALECTOTYPE| Staphylinus| cinerarius Erich-
(synonym of C. m. maxillosus); (unavailable: infra- son, 1839| designated by| D. J. Clarke 2008’ (in
subspecific name, Article 45.6.4). ZMHB).
Creophilus maxillosus var. subfasciatus Sharp.

‘Syntype’. 씹, ‘[circular with red border] Type/ Japan.|
G Lewis./ Sharp Coll.| 1905–313./ C. maxillosus var.|
subfasciatus| Type D.S./ [yellow circular] Japan/
FMNH-INS 0000 016 793/ [red] SYNTYPE| Creophi-
lus maxillosus| var. subfasciatus Sharp, 1874| teste
D. J. Clarke 2008’ (in BMNH). Specimen pinned to
card, aedeagus and genital segment glued to card.
Creophilus maxillosus var. medialis Sharp.
‘Syntype’. 씹, ‘[circular with red border] Type/ Japan.|
G. Lewis./ Sharp Coll.| 1905–313./ C. maxillosus
var.| medialis| Type D.S./ [yellow circular] Japan/
FMNH-INS 0000 016 794/ [red] SYNTYPE| Creophi-
lus maxillosus| var. medialis Sharp, 1874| teste D. J.
Clarke 2008’ (in BMNH). Specimen pinned to card,
aedeagus glued to card.
Creophilus maxillosus var. imbecillus Sharp. ‘Syn-
types’ (2). 씹, ‘[circular with red border] Type/
Japan.| G. Lewis./ Sharp Coll.| 1905–313./ C. max-
illosus var.| imbecillus| Type D.S./ [yellow circular]
Japan/ FMNH-INS 0000 016 792/ [red] SYNTYPE|
Creophilus maxillosus| var. imbecillus Sharp, 1874|
teste D. J. Clarke 2008’ (in BMNH) – front left
tarsal claw and right hind tibia missing; 씹,
‘Japan.| G. Lewis./ Sharp Coll.| 1905–313./ C. max-
illosus var.| imbecillus| mihi D.S./ FMNH-INS
0000 019 494/ [red] SYNTYPE| Creophilus maxillo-
sus| var. subfasciatus Sharp, 1874| teste D. J. Figure 14. Ventral vestiture patterning of A, C. maxillo-
Clarke 2008’ (in BMNH). The two specimens here sus maxillosus and B, C. m. villosus. Scale bar = 10 mm.
identified as ‘syntypes’ each possess a label written
in Sharp’s handwriting and initialled with ‘D.S’.
basolateral pronotal denticle of large males distinctly
Both match his description in having yellowish
pointed (Fig. 2G, bpd); appendix of internal sac
pubescence and abrasion.
pointed (Fig. 4B, app).
Creophilus maxillosus ssp. canariensis Bernhauer.
Lectotype (here designated). 씹, ‘Fuerteventura/
Canar. Ins.| Polatzek./ maxillosus L.| det. Bernh./ Measurements: (N = 10씹, 10씸) Forebody length: 씹
[Bernhauer’s handwriting] ssp.canariensis| Brnh. 5.7–10.3, 씸 6.3–8.7 mm. See supporting Table S4 for
Typus./ Chicago NH Mus| M. Bernhauer| Collection/ comparison of ranges of male and female ratios.
FMNH-INS 0000 017 792/ [red] LECTOTYPE| Creo- HW/HL = 1.39–1.70; EYL/HL = 0.46–0.71; HL1/
philus maxillosus ssp.| canariensis Bernhauer, 1908| HL2 = 1.43–3.50; ML/HL = 0.94–1.50; ML1/
designated by| D. J. Clarke 2008’ (in FMNH). ML2 = 0.60–0.92; PW/PL = 1.07–1.24; HL/PL = 0.65–
Creophilus sikkimensis Wendeler. Holotype (by 0.83; HW/PW = 0.86–1.10; PL/ESL = 1.31–1.50.
monotypy). 씹, ‘씹/ Sikkim/ Erworben von| Alexander
Heyne| Berlin – Wilmersd./ Creophilus| sikkimensis Variation: Creophilus maxillosus maxillosus exhibits
n.sp.| Wendeler det./ [pink] Holotypus/ Zool. Mus.| variability primarily in vestiture patterns, but with
Berlin/ Creophilus| maxillosus L.| det. Schillham- less extensive patterns being a subset of the
mer, 2003/ FMNH-INS 0000 016 789/ [red] HOLO- maximum pattern that is seen in Egyptian and
TYPE| Creophilus| sikkimensis Wendeler, 1927| Canary Islands specimens. Rare forms of C. m. max-
teste D. J. Clarke 2008’ (in ZMHB). illosus, such as the C. ciliaris holotype, are charac-
terized by a more evenly extensive golden-brown or
Diagnosis: With characters of the maxillosus-group; reddish vestiture.
vestiture on hind margins of head, anterior pronotal
declivities, and pterothorax (Fig. 14A) mostly black; Remarks: In erecting his three new names (var. sub-
vestiture on dorsal surface of abdomen white, fasciatus, var. medialis, var. imbecillus), it is clear
arranged into definite pattern, concentrated on terg- that Sharp (1874) was referring to infrasubspecific
ites V and VI, with VII–X usually partly whitish; variation: ‘A considerable number of a species of Creo-
764 D. J. CLARKE
philus, brought by Mr Lewis from Japan, should, I maxillosus s.l.); Smetana, 2004: 672 (synonym of C.
think, be referred to as a variety of C. maxillosus.’ He m. maxillosus).
then described variation within this series, ‘The speci- Staphylinus arcticus Erichson, 1839: 348. Type
mens, however, are very discrepant among them- locality: ‘Kamtschatka’; Ganglbauer, 1895: 415
selves . . .’, and suggested the three names for the (synonym of C. maxillosus s.l.); Herman, 2001b: 3323
forms he recognized. His concluding sentence ‘. . . I (synonym of C. maxillosus s.l.); Navarrete-Heredia
think it not unlikely it will be ultimately found that et al., 2002: 322 (synonym of C. maxillosus s.l.);
C. maxillosus, cinerarius, arcticus, and villosus Smetana, 2004: 671 (synonym of C. m. maxillosus).
cannot be maintained as distinct species, but must be Creophilus acuticollis Bernhauer, 1910: 377.
looked on as geographical races of one and the same Type locality: ‘Mexiko’; Herman, 2001b: 3315;
species’ clearly indicates he meant his names to Navarrete-Heredia et al., 2002: 322 (synonym of C.
describe local (Japanese) variation within one of these maxillosus s.l.).
‘races’, i.e. C. m. maxillosus. To my knowledge, these Creophilus maxillosus villosus ab. ciliaroides
names have not been adopted as the valid names of a Hatch, 1938: 149. Type locality: ‘Soap Lake, Washing-
species or subspecies before 1985, nor treated as ton’; Herman, 2001b: 3324 (synonym of C. maxillosus
senior homonyms, and in accordance with Article s.l.; as Creophilus maxillosus villosus var. ciliaroides);
45.6.4 (ICZN, 1999), it is concluded that var. Navarrete-Heredia et al., 2002: 322 (synonym of C.
subfasciatus, var. medialis, and var. imbecillus are maxillosus s.l.); Smetana, 2004: 672 (synonym of C.
infrasubspecific names and therefore unavailable. m. maxillosus) (unavailable: infrasubspecific name,
Examination of the series of C. maxillosus specimens Articles 45.6.1, 45.6.2).
seen by Sharp reveals that there are no consistent
differences between them and the typical (European) Type material: Staphylinus villosus Gravenhorst. Lec-
form of C. maxillosus. totype (here designated). 씹, ‘5877/ [red] Typus/ [red]
LECTOTYPE| Staphylinus| villosus Gr.| designated
by| A. Newton 1988/ [green] villosus| Gr.| Am. Spt./
Distribution: Widespread in the continental Palearc- Zool. Mus.| Berlin/ FMNH-INS 0000 016 795/ [red]
tic, and on most Atlantic islands. Adventive in LECTOTYPE| Staphylinus| villosus Gravenhorst,
southern South America (Chile, Argentina, Peru: 1802| designated by| D. J. Clarke 2008’ (in ZMHB).
Fig. 24). Asenjo & Clarke (2007) recorded C. Paralectotypes (2). 씹, ‘[green] Am. sept.| Nr. 5877/
maxillosus s.l. from Peru. These specimens all [red] Typus/ Zool. Mus.| Berlin/ [yellow] PARALEC-
belong to C. m. maxillosus, and have the vestiture TOTYPE| Staphylinus| villosus Gr.| designated by|
patterning and variation typical of that subspecies. A. Newton 1988/ FMNH-INS 0000 016 796/ [yellow]
Creophilus maxillosus maxillosus is also found in PARALECTOTYPE| Staphylinus| villosus Graven-
western Alaska and the north eastern US (Newton horst, 1802| designated by| D. J. Clarke 2008’ (in
et al., 2000; Fig. 15, filled circles), possibly due to ZMHB); 씹, same label data, and ‘FMNH-INS
human-assisted dispersal. Smetana (2004) recently 0000 016 797/ [yellow] PARALECTOTYPE| Staphyli-
gave a comprehensive summary of the distribution nus| villosus Gravenhorst, 1802| designated by|
of this subspecies. D. J. Clarke 2008’ (in ZMHB). Newton’s lectotype
designation has not been published (A. F. Newton,
pers. comm.).
Creophilus Fasciatus Laporte de Castelnau.
1B. CREOPHILUS MAXILLOSUS VILLOSUS Syntype. 씸, ‘amer. bor/ fasciatus Lap| type/ Kansas/
(GRAVENHORST) Maryland/ whitewood| Canada/ Coll. et det. A.
(FIGS 4C, 5A, 9C–F, L, M, 10B, 13H, Fauvel| CREOPHILUS| maxillosus| var. villosus
14B, 15, 19B) Grav| R.I.Sc.N.B. 17.479/ [red] TYPE?/ FMNH-INS
Staphylinus villosus Gravenhorst, 1802: 160. Type 0000 016 784/ [red] SYNTYPE| Creophilus| Fascia-
locality: ‘Am. septentr.’; Sharp, 1884: 370 (variety of tus Laporte, 1835| teste D. J. Clarke 2008’ (in
C. maxillosus s.l.); Herman, 2001b: 3322 (synonym of IRSNB). Because of uncertainty surrounding Fauvel’s
C. maxillosus s.l.); Navarrete-Heredia et al., 2002: multiple locality labelling system, I refrain from des-
322 (synonym of C. maxillosus s.l.; as nom. praeocc., ignating this putative syntype as the lectotype (see
nec DeGeer, 1774 [incorrect]). Material and Methods),
Creophilus Fasciatus Laporte de Castelnau, 1835: Staphylinus arcticus Erichson. Holotype (by
111. Type locality: ‘Amérique du nord’; Herman, monotypy). 씸, ‘5879/ [orange] Type/ [yellow] arcticus|
2001b: 3323 (preoccupied by Staphylinus fasci- Er.| Kamtschatka v Cham/ [white] Hist.-Coll.
atus Füessly, 1775, synonym of C. maxillosus s.l.); (Coleoptera)| Nr. 5879| Staphylinus arcticus|
Navarrete-Heredia et al., 2002: 322 (synonym of C. Erichs.| Kamtschatka, Chamisso| Zool. Mus. Berlin/
Figure 15. Distribution of all georeferenced specimens of C. maxillosus villosus (white circles) and C. m. maxillosus
(black circles) in North America. Creophilus m. villosus is also recorded from Guatemala, Panama, and Chile (no locality).
The holotype of C. arcticus, a synonym of C. m. villosus, is from Russia: Kamchatka.
[red] HOLOTYPUS| Staphylinus| arcticus Erichson, ites IV and V, with tergites III and VI–X usually
1839| labeled by MNHUB 2004/ FMNH-INS completely black; basolateral pronotal denticle of
0000 016 785/ [red] HOLOTYPE| Staphylinus| arcti- large males rounded or otherwise indistinct; appendix
cus Erichson, 1839| teste D. J. Clarke 2008’ (in of internal sac slightly to distinctly flared laterally
ZMHB). (Figs 4C, 10B, app).
Creophilus acuticollis Bernhauer. Holotype (by
monotypy). 씹, ‘[pink] Hoege| Mexico/ [peach] Measurements: (N = 10씹, 10씸). Forebody length: 씹
villosus/ acuticollis| Brnh. Typus/ [red] HOLOTYPE| 5.7–10.0 mm, 씸 5.9–8.4 mm. See supporting Table S4
Creophilus| acuticollis| Bernh./ FMNH-INS for comparison of ranges of male and female ratios.
0000 016 790/ Creophilus| maxillosus L.| Newton HW/HL = 1.39–1.63; EYL/HL = 0.47–0.63; HL1/HL2 =
det. 1974/ [red] HOLOTYPE| Creophilus| acuticollis 1.46–3.60; ML/HL = 0.94–1.41; ML1/ML2 = 0.71–0.91;
Bernhauer, 1910| teste D. J. Clarke 2008’ (in FMNH). PW/PL = 1.08–1.23; HL/PL = 0.64–0.84; HW/PW =
Creophilus maxillosus villosus ab. ciliaroides 0.89–1.07; PL/ESL = 1.23–1.50.
Hatch. ‘Holotype’ (original designation). 씸, ‘Soap L.,
WASH.| April 19, 1935| M. H. Hatch/ [red] TYPE| Distribution (Fig. 15, open circles): Widespread in
Creophilus| maxillosus| Ssp. villosus| ab. ciliaroi- North America; Aleutian Islands; Hawaii; extending
des| 1938-M.H. Hatch/ Pub. as| Staphylinus| max- south into Mexico and northern Central America, and
illosus| villosus| ab. ciliaroides| Hatch, 1938/ in Cuba, Jamaica, and Dominican Republic in the
FMNH-INS 0000 019 495/ [red] HOLOTYPE| Creo- West Indies. Because the holotype of C. arcticus is
philus maxillosus villosus| ab. ciliaroides Hatch, from Kamtschatka Peninsula the range is extended to
1938| teste D. J. Clarke 2008’. Missing right protar- that region of the Palearctic; it is likely that both
sal claw, right metatarsus adhered to first label (in subspecies are distributed outside their native ranges
USNM). ‘Paratypes’ (4) (in OSAC; not seen). because of human-assisted dispersal.
Diagnosis: With characters of the maxillosus-group; Remarks: The C. fasciatus syntype listed above exhib-
vestiture on hind margins of head, anterior pronotal its typical C. m. villosus coloration; the C. arcticus
declivities, and pterothorax (Fig. 14B) mostly white; holotype is also typical of C. m. villosus in having
vestiture on dorsal surface of abdomen white whitish setae on the anterior pronotal angles, hind
arranged into definite pattern concentrated on terg- margins of head, and concentrated on tergites IV–V.
766 D. J. CLARKE
The setae of the pterothorax, however, are much probably in Paris Museum’. Fauvel did not indicate
darker. The holotype of C. acuticollis is an aberrant the number of specimens examined.
specimen of C. m. villosus; the prothorax is strongly
constricted at the middle, and the pro- and mesocoxae Other material examined: 170 specimens. See sup-
are conspicuously deformed. It is clear from the name porting information, Appendix S1.
(C. m. villosus ab. ciliaroides) and from Hatch’s (1938:
148–149) comments that he was describing an Diagnosis: With characters of the maxillosus-group;
infrasubspecific taxon: ‘Since it is not desirable integument and elytra uniformly black; antenna
to apply the same varietal name [ciliaris] in two distinctly clavate; antennomere 11 convex apically
distinct subspecies [maxillosus and vilosus], I desig- (Fig. 16B); basolateral pronotal impressions marked
nate this form ab. ciliaroides nov. (type and four by deep punctule (Fig. 16G); basolateral pronotal
paratypes).’ In accordance with Article 45.6.4 (ICZN, macroseta absent; elytral vestiture with indistinct
1999), the name ciliaroides applies to an infrasubspe- whitish- or silvery-grey patch (Figs 1G, 16H); tergal
cific category and is therefore unavailable. chaetotaxic formula 6-6-6-0(2)-4-6.
Description: Measurements (N = 10씹, 10씸). Forebody

2. CREOPHILUS INCANUS (KLUG)
length: 씹 5.8–11.0 mm, 씸 6.1–8.0 mm. See surpport-
(FIGS 1G, 2K, 4F, 16, 17) ing Table S4 for comparison of ranges of male and
Staphylinus (Emus) incanus Klug, 1834: 137. Type female ratios. Head. Head weakly trapezoidal and
locality: Madagascar. widest posteriorly to subrectangular, particularly in
Staphylinus incanus; Erichson, 1839: 350; Vinson, females; HW/HL = 1.43–1.70; basal margins sparsely
1956: 51; Vinson, 1967: 320. setose; dorsal punctation sparse; eyes large (EYL/
Staphylinus (Creophilus) Coquerelii Fauvel, 1866: HL = 0.48–0.65), dorsolateral in large males, more
317. Type locality: Réunion: Bourbon; 1867: 67; 1868: lateral in females and smaller males, not protruding,
71 (same as 1867 paper); Vinson, 1956: 51 (synonym lateral margins of head visible in dorsal view only in
of C. incanus); Lecoq, 1990: 206 (as coquereli, large males, obscured by eye in smaller specimens;
synonym of C. incanus). HL1/HL2 much greater in females than males
Creophilus incanus; Fauvel, 1875: 55; Alluaud, (씹 = 1.21–2.40, 씸 = 2.60–3.25); antennae as in
1900: 87; Bernhauer & Schubert, 1914: 398; Figure 16B, distinctly clavate; antennomeres 1–6
Lecoq, 1990: 206, 210, fig. 39; Herman, 2001b: brownish-black, 7–11 greyish-black, 9–10 narrowed
3317. anteriorly, thickened posteriorly; apex of antenno-
Creocephalus incanus; Fairmaire, 1893: 523 (error mere 11 convex medially; each pair of apical setae
for Creophilus, see Blackwelder, 1952: 111). moderately widely separated, one on each side of
Creophilus Coquereli; Alluaud, 1900: 88; Bernhauer apex; mandibles as in Figure 16A, longer than head
& Schubert, 1914: 398 (synonym of C. incanus). in large males, subequal to head in females and small
males (ML/HL 씹 = 0.88–1.48, 씸 = 0.94–1.06), T1
Type material: Staphylinus (Emus) incanus Klug. Lec- largest, T3 hardly visible in dorsal view. Thorax and
totype (here designated). 씹, ‘[orange] Type/ incanus abdomen. Pronotum (Fig. 16G) slightly transverse
Kl./ 5884/ [green] incanus| Kl.| Madag Goud./ [blue] (PW/PL = 1.07–1.24), moderately to distinctly nar-
Hist.-Coll. (Coleoptera)| Nr. 5884| Staphylinus rowed posteriorly; PL 1.24–1.56 ¥ ESL; basolateral
incanus Kl.| Madagasc., Goudot| Zool. Mus. Berlin/ margins very shallowly emarginate, hind angles dis-
[red] SYNTYPUS| Staphylinus| incanus Klug, 1832| tinct, obtuse; disc surface faintly sinuous laterally,
labeled by MNHUB 2004/ FMNH-INS 0000 016 781/ with sparse, black, peripheral setae and very sparse
[red] LECTOTYPE| Staphylinus (Emus)| incanus short vestiture on lower anterolateral declivities;
Klug, 1832| designated by| D. J. Clarke 2008’ (in basolateral impressions sparsely setose, accentuated
ZMHB). Specimen missing left metatarsus and right by deep punctule; scutellum with uniformly black
metatarsal claws. Paralectotype. 씸, ‘[orange] Type/ vestiture; elytra uniformly black to brownish-black,
[blue] Hist.-Coll. (Coleoptera)| Nr. 5884| Staphylinus vestiture sparse, with poorly delimited central white
incanus Kl.| Madagasc., Goudot| Zool. Mus. Berlin/ patch (Fig. 16H); wings fully developed, clear
[red] SYNTYPUS Staphylinus| incanus Klug, 1832| yellowish-brown, without black spot in medial field
labeled by MNHUB 2004/ FMNH-INS 0000 016 782/ between MP3 and MP4 veins; vestiture of pterothorax
[yellow] PARALECTOTYPE| Staphylinus (Emus)| mostly whitish, that of legs mostly black; dorsal
incanus Klug, 1832| designated by| D. J. Clarke abdominal vestiture mostly black but with variable
2008’ (in ZMHB). maculation of whitish vestiture concentrated on terg-
Staphylinus (Creophilus) Coquerelii Fauvel. Syn- ites and parasclerites III–V (Fig. 1G); vestiture of
types (?; not seen). Vinson (1956) notes ‘type: Réunion, sternites III–VI mostly whitish, interrupted by more
Figure 16. Creophilus incanus, details of morphology (setae of mandibles, antenna, and pronotum omitted). A, man-
dibles, dorsal (bases omitted); B, right antenna, dorsal (only apical setae shown; anterior is to the left); C, median lobe
apex, ventral; D, paramere, dorsal; E, apical part of median lobe with fully everted internal sac, right lateral; F, aedeagus
with details of inverted internal sac, right lateral (parameral setae omitted); G, pronotum (small male); H, right elytron
(only whitish setae shown); I, female internal genitalia, ventral (surface of vf with minute teeth as shown for vp of C.
maxillosus villosus in Fig. 9M); J, ventral sclerite of internal sac, ventral (lower) and apical (upper). Abbreviations: T1,
T2, T3, mandibular teeth. Additional abbreviations as in Figure 3 (male) and Figure 5 (female). Scale bars = 1 mm.
or less circular black patches laterally, vestiture on (Fig. 16I, vf) with embedded minute tooth-like scales
sternites VII–IX mostly black; whitish abdominal ves- (as in Fig. 9M). Chaetotaxy. Basolateral pronotal mac-
titure frequently with bluish-grey tinge; abdominal rosetae absent; elytral discal series with 3–4 macrose-
tergite VII with well-developed palisade fringe. Male tae; tergal chaetotaxic formula = 6-6-6-0(2)-4-6, inner
genitalia. Aedeagus as in Figure 16F; median lobe lateral macrosetae absent from tergite VII, tergite VI
apex narrowly subacuminate (Fig. 16C); paired apico- usually without macrosetae, outer lateral macrosetae
lateral sclerites (as) articulated to internal edge of occasionally present.
median orifice, not fused to it. Paramere as in
Figure 16D. Internal sac inverted as in Figure 16F,
everted as in Figure 16E; ventral sclerite (vs) deeply Variation: Five specimens from Réunion and three
notched apically (Fig. 16J); ventral process (vp) short, from Madagascar have mostly black vestiture
with ventromedian spiculose strip (vr) shorter than on elytra, tergites, and ventral side, with only
ventral sclerite (vs). Female internal genitalia. Inter- paired groups of white setae in marginal fringes
nal female genitalia as in Figure 16I; vaginal plate of tergites III–VI. Aedeagi of males from Réunion
(vp) broadly rounded to truncate posteriorly, with and Mauritius were identical to those from
median sclerotized strip (ss); surface of vaginal fold Madagascar.
768 D. J. CLARKE
data). Phenology: February–December. Vinson

(1956) records one specimen ‘on decaying Clemora
beetles’ (Scarabaeidae: junior synonym of Phyllo-
phaga Harris), another ‘trapped in an Aristolochia
flower’. He notes the species is ‘most probably a
predator of dipterous larvae in carrion’.
Remarks: A female specimen from Réunion in Bern-

hauer’s collection (FMNH) bears the label ‘nigerrimus
Typus’ in Bernhauer’s handwriting. Another ‘nigerri-
mus’ specimen (syntype?) is possibly in NMW (A. F.
Newton, pers. comm.). Further collecting on Réunion
may reveal subspecific differences from Madagascan
specimens.
3. CREOPHILUS GALAPAGENSIS CLARKE SP. NOV.

(FIGS 1H, 4D, 18, 19A, 20)
Type material: Holotype. 씹, ‘ECUADOR: Galápagos:|
Isla Isabela, Volcán| Alcedo (NE rim), 1100 m,|
0 °24′S, 91 °07′W,| 21–25.vi.1991/ shrub forest:
FMHD#91-|203, Peck#91–246, carrion| traps, S.
Peck| FIELD MUS. NAT. HIST./ FMNH-INS
0000 019 454/ [red] HOLOTYPE| Creophilus| gal-
apagensis Clarke sp. nov.| designated by| D.J.
Clarke 2010’ (in FMNH). The aedeagus and genital
segments are dissected and stored in glycerin in a
genitalia vial mounted with specimen. Paratypes
(1069): all pinned specimens and alcohol lots with
label ‘[yellow] PARATYPE| Creophilus| galapagensis
Clarke sp. nov.| designated by| D.J. Clarke 2010’. 3
씸, with labels ‘ISLAS GALAPAGOS| ISABELA|
12–11/85| lg. S. Abedrabbo/ ex.| AVE MUERTA [dead
bird]/ 5/ Creophilus| maxillosus| (Linnaeus)| det.
Newton 2003’: ‘Creophilus| maxillosus| DET: S.Peck,
2002/ FMNH-INS 0000 019 470’; ‘FMNH-INS
0000 019 471’; ‘FMNH-INS 0000 019 472’ (in FMNH).
3 specimens, ‘Isabela| Cerro Azul [0 °54′S 91 °25′W]|
Figure 17. Distribution of all georeferenced specimens of
C. incanus in Madagascar and Mayotte (circles). This
[underside of label]SA 308–309/ Islas Galapagos|
species is also known from Réunion and Mauritius islands. lg.H.Franz,V.-VI, 1975’: 2씹, ‘FMNH-INS
0000 017 435’; ‘FMNH-INS 0000 017 436’; 1씸,
‘FMNH-INS 0000 017 437’ (in NMW). 2 specimens,
Comparison: Creophilus incanus resembles C. gal- ‘Albermarle I. [Isla Isabela]| Galapagos Is./ Cobes|
apagensis and C. maxillosus villosus in having the Settlement| IV.24.06/ Coll. By F.X. Williams’: 씹,
whitish dorsal abdominal vestiture concentrated on ‘FMNH-INS 0000 017 808’; 씸, ‘[in pencil] S.A.| 31/
tergites IV–V. It is distinguished from both most FMNH-INS 0000 017 809’ (in CAS). 133 specimens,
easily by the convex apex of the terminal antenno- ‘ECUADOR: Galápagos:| Isla Isabela, Santo| Tomás,
mere and the posteriorly thickened preceding two 350 m,| 0 °51′S, 91 °02′W,| 6–9.vii.1985, mixed
antennomeres (Fig. 16B). forest;| FMHD#85–1037,| Peck#85–214, carrion|
traps, S. & J. Peck| FIELD MUS. NAT. HIST.’: 58씹
Distribution (Fig. 17): Madagascar, Mauritius, in 70% alcohol, with lot ID number ‘FMNH-INS
Mayotte (Alluaud, 1900), and Réunion. 0000 017 800’; 42씸 in 70% alcohol, with lot ID
number ‘FMNH-INS 0000 017 801’; 19씹 pinned, 8
Biology and ecology: Rare in collections. Two speci- with aedeagi dissected, FMNH-INS 0000 017 810–
mens were taken from cow dung. Habitat: unknown. 828; 5씹 pinned, 4 with aedeagi dissected, FMNH-INS
Altitude: 800 m (only one specimen with elevation 0000 017 625–629; 9씸, [FMNH-INS 0000 017 616–
618; 829–834] (in FMNH). 1씹, ‘ECU.Galap.Isabela| Volcán Cerro| Azul, 7 km NE Caleta| Iguana,
Sto. Tomas,300–350 m/ 9–12.III.89, [Peck#]89–126| 700 m,| 0 °56′S, 91 °26′W./ 20–25.V.1991, pampas;|
agric.z.carrion tp.| B.J Sinclair/ Creophilus| maxil- FMHD#91–181, Peck#91-|155, carrion trap, S. Peck|
losus| (Linnaeus)| det. Newton 2003/ FMNH-INS FIELD MUS. NAT. HIST.’: 41씹 in 70% alcohol, with
0000 019 473’ (in FMNH). 1 씹, aedeagus dissected on lot ID number ‘FMNH-INS 0000 017 798’; 43씸
card, ‘Tagos [Tagus] Cove [0 °16′S 91 °22′W]| Alber- in 70% alcohol, with lot ID number ‘FMNH-INS
marle I. [Isla Isabela]| Galapagos Is./ IV-1–06/ Coll. 0000 017 799’; 2씹 pinned, ‘FMNH-INS 0000 017 794’;
by| F.X. Williams/ FMNH-INS 0000 017 807’ [para- ‘FMNH-INS 0000 017 795’; 2씸 pinned, ‘FMNH-INS
sitic worm in genitalia vial] (in CAS). 3 씹, all with 0000 017 796–797’; 1씸 pinned, with measurements
aedeagi dissected, ‘Villamil [0 °56′S 91 °1′W]| Alber- taken, ‘FMNH-INS 0000 017 619’ (in FMNH). 6
marle I. [Isla Isabela]| Galapagos Is./VIII-22– specimens, ‘Isabela| Umg. Sta. Tomas [in area of
06[1906]/ Coll. by| F.X. Williams’: ‘FMNH-INS Santo Tomás]| [underside of label]SA 299/ Islas Gal-
0000 017 804/ Creophilus| villosus| (Grav)’; ‘FMNH- apagos| lg.H.Franz,V.-VI, 1975’: 3씹, [FMNH-INS
INS 0000 017 805/ [in pencil] S.A. 44.’; ‘FMNH-INS 0000 017 438–440]; 3씸, ‘FMNH-INS 0000 017 441/
0000 017 806/ [in pencil] S.A.| 46.’ (in CAS). 5 speci- Creophilus| villosus Grav.| det.H.Franz’; ‘FMNH-
mens, ‘ECUADOR: Galápagos I.| Isla Isabela, INS 0000 017 442–443’ (in NMW).
Volcán| Alcedo, crater rim [0 °25′S, 91 °6′W], 1000|
m, 2.IV.1996, hand coll-| ecting, S. Peck 96–84| Diagnosis: With characters of the maxillosus-group;
FIELD MUS. NAT. HIST.’, and with det. label integument and elytra uniformly pitchy black; elytral
‘Creophilus| maxillosus| (Linnaeus)| det. Newton vestiture entirely black (Fig. 1H); elytra rugose;
1998’: 씹, with aedeagus dissected ‘FMNH-INS abdominal microsculpture imbricate (Fig. 19A); tergal
0000 017 416’; 4씸 [measurements taken, FMNH-INS chaetotaxic formula 2-2-2-2-2-4.
0000 017 417–420] (in FMNH). 779 specimens, same
data as holotype: 349씹 in 70% alcohol, with lot Description: Measurements (N = 10씹, 10씸). Forebody
ID number ‘FMNH-INS 0000 017 802’; 381씸 in length: 씹 5.3–8.7 mm, 씸 5.4–7.5 mm. See supporting
70% alcohol, with lot ID number ‘FMNH-INS Table S4 for comparison of ranges of male and female
0000 017 803’; 5씹 pinned [measurements taken, ratios. Head. Head subtrapezoidal to subrectangular,
FMNH-INS 0000 017 620–624]; 23씹 pinned, [FMNH- widest posteriorly; large males with slight hind
INS 0000 017 857–879]; 1씹 pinned, [FMNH-INS angles evident; HW/HL = 1.38–1.54; basal margins
0000 019 440]; 13씸 pinned, [FMNH-INS almost asetose; dorsal punctation dense and coarse;
0000 019 441–453]; 5씹 pinned, with aedeagi dis- eyes large (EYL/HL = 0.50–0.67), dorsolateral in large
sected, [FMNH-INS 0000 019 455–459]; 1씹 pinned, males, more lateral in females and smaller males,
‘FMNH-INS 0000 019 460’ (in FMNH); 1씹, 1씸 prominent and moderately protruding, lateral
pinned, deposited in each of AMNH, BMNH, BPBM, margins of head obscured by eye in dorsal view;
CAS, CDRS, CMNC, IRSNB, MCZ, NMW, QCAZ, HL1/HL2 not distinctly greater in females than
SEMC, and USNM. 1씹, same locality and habitat males (씹 = 1.45–3.67, 씸 = 3.00–4.00); antennae as in
data as holotype, ‘. . . pitt &| bottle tps under| Figure 18B, antennomeres 1–6 black, 7–11 greyish-
shrubs, SPeck, 91–245/ Creophilus| maxillosus| (Lin- black; apex of antennomere 11 moderately to deeply
naeus)| det. Newton 2003/ FMNH-INS 0000 019 475’ emarginate medially; each pair of apical setae widely
(in FMNH). 22 specimens, ‘ECUADOR: Galápagos:| separated by apical emargination, one on each side of
Isla Isabela, Volcán| Alcedo (NE slope), 850 m,| apex; mandibles as in Figure 18A, longer than head
0 °22′S, 91 °06′W,| 20–25.vi.1991/ open forest: in large males, shorter or subequal to head in females
FMHD#91-|201, Peck#91–243, carrion| trap, S. (ML/HL 씹 = 0.87–1.30, 씸 = 0.81–1.00), T1 larger
Peck| FIELD MUS. NAT. HIST.’: 10씹, [FMNH-INS than T2 and T3. Thorax and abdomen. Pronotum
0000 017 835–844]; 12씸, [FMNH-INS 0000 017 845– (Fig. 18F) slightly transverse (PW/PL = 1.07–1.24),
856] (in FMNH). 1씹, same locality and habitat data, moderately to strongly narrowed posteriorly; PL 1.24–
‘. . . night| colln, S. Peck, 91–244/ Creophilus| max- 1.44 ¥ ESL; basolateral margins faintly incised, hind
illosus| (Linnaeus)| det. Newton 2003/ FMNH-INS angles indistinct; basolateral denticle of large males
0000 019 474’ (in FMNH). 14 specimens, ‘ECUADOR: indistinct; with irregularly scattered peripheral setae
Galápagos I.| Isla Isabela| Cerro Azul,| 5 km NE and sparse short setae on lower anterolateral declivi-
Caleta Iguana [0 °56′S, 91 °27′W],| 400 m, pampas;/ ties; basolateral impressions asetose, asperate; scutel-
20–25.V.1991, carrion| trap, S. Peck 91–154| FIELD lum imbricate with black vestiture; elytra uniformly
MUS. NAT. HIST.’: 11씹, [FMNH-INS 0000 017 421– black, rugosely sculptured, sparsely covered with very
431]; 3씸, [FMNH-INS 0000 017 432–434; -433 and fine uniformly black vestiture (Fig. 1H); glabrous
-434 with measurements taken] (in FMNH). 89 humeral region large, impunctate; wings fully devel-
specimens, ‘ECUADOR: Galápagos I.| Isla Isabela, oped, black, but without distinct black spot in medial
770 D. J. CLARKE
Figure 18. Creophilus galapagensis, details of morphology (A, C, E, F, I = holotype, small male); setae of mandibles,
antenna, and pronotum omitted). A, mandibles, dorsal (bases omitted); B, right antenna, dorsal (only apical setae shown);
C, median lobe apex, ventral; D, ventral sclerite of internal sac, ventral (lower) and apical (upper); E, aedeagus with
details of inverted internal sac, right lateral (parameral setae omitted); F, pronotum; G, female internal genitalia, ventral;
H, apical part of median lobe with fully everted internal sac, right lateral; I, paramere, dorsal. Abbreviations: T1, T2, T3,
mandibular teeth. Additional abbreviations as in Figure 3 (male) and Figure 5 (female). Scale bars: D = 0.1 mm;
rest = 1 mm.
field between MP3 and MP4 veins; ventral pterotho- fused to it. Paramere as in Figure 18I. Internal sac
racic vestiture golden brown, vestiture of legs black; inverted as in Figure 18E, everted as in Figure 18H;
abdomen with dull reflection due to characteristically ventral process (vp) with ventromedian spiculose
imbricate microsculpture (Fig. 19A), with short strip (vr) > 2¥ length of ventral sclerite (vs); membra-
appressed and moderately dense vestiture; parascler- nous sheath of copulatory piece (cp) reduced, with
ites of abdominal segment II completely fused most of apical portion of cp exposed; appendix flared
(without apical notch); dorsal abdominal vestiture laterally (Fig. 4D, app). Female internal genitalia.
mostly black, with golden brown maculation mostly Female genitalia as in Figure 18G; vaginal plate (vp)
on tergites IV–V (Fig. 1H); ventral abdominal vesti- broadly bilobed posteriorly, with median sclerotized
ture golden brown on sternites III–VIII, with more strip (ss); vaginal fold completely membranous, bursa
black on VII and VIII; tergite VII with well-developed copulatrix (bcx) distinctly longer than vaginal plate;
palisade fringe. Male genitalia. Aedeagus as in spermatheca (sp) sausage-like, connected to bcx via
Figure 18E; median lobe apex short, narrowly sub- short duct. Chaetotaxy. Basiantennal and basolateral
acute (Fig. 18C); paired apicolateral sclerites (as) pronotal macrosetae absent or vestigial, rarely
articulated to internal edge of median orifice, not present as pair; basisternal pair absent; ps1 absent,
Figure 19. Scanning electron micrographs of abdominal tergal microsculpture. A, Creophilus galapagensis, imbricate; B,
C. maxillosus villosus (Gravenhorst), transversely strigulate. Scale bars = 0.1 mm.
humeral macroseta usually absent, if present then

rarely on both elytra; elytral discal series with 3–4
macrosetae; posterior elytral patch variable, baso-
humeral macrosetae usually absent, if present only
one, and rarely on both elytra, posterior epipleural
macroseta present or absent; tergal chaetotaxic
formula = 2-2-2-2-2-4 (unique in Creophilus), medial
macrosetae absent on tergites III–VII; inner lateral
macrosetae absent on tergites III–VIII; second gono-
coxal macroseta usually absent, if present rarely on
both gonocoxites.
Variation: The extent of abdominal vestiture pattern-

ing varies little, with tergites IV and V consistently
with densest golden brown vestiture and other terg-
ites with either sparse golden brown patches or
entirely black vestiture. Chaetotaxy of head, prono-
tum, and elytra is unusually variable.
Comparison: Creophilus galapagensis superficially

resembles C. maxillosus villosus and C. incanus due
to golden-brown abdominal vestiture concentrated on
tergites IV and V, and golden-brown pterothoracic
vestiture. The entirely black elytral vestiture, imbri-
cate abdominal microsculpture (Fig. 19A), and
reduced abdominal chaetotaxy (all unique characters
within the C. maxillosus-group) will consistently dis-
tinguish C. galapagensis from these two species. Creo-
philus galapagensis is distinctly smaller than its
congeners; males attain a maximum size only two-
thirds that of C. maxillosus males.
Figure 20. Distribution of all georeferenced specimens of
C. galapagensis on Isla Isabela in the Galapagos Islands
Distribution (Fig. 20): Galápagos Islands: Isla (circles). The species is also known from Fernandina, San-
Isabela; Fernandina and Santiago (Newton & Peck, tiago and San Cristóbal Islands.
2006, as C. maxillosus, see below); San Cristóbal
(Darwin’s specimens, see below).
772 D. J. CLARKE
Biology and ecology: Most collection records are from Emus insularis Fauvel, 1879: 94. Type locality:
carrion traps. Other records are from under a dead ‘Batcian’; new synonymy.
bird (Waterhouse, 1845) and from a decaying duck Creophilus insularis; Bernhauer & Schubert, 1914:
(Franz, 1985). Habitat: agricultural areas, mixed 399; Bernhauer, 1926: 349; Scheerpeltz, 1933: 1412;
forest, shrub forest, open forest, and pampas grass- Herman, 2001b: 3326.
land. Altitude: 325–1100 m. Phenology: March– Creophilus violaceus; Bernhauer & Schubert, 1914:
November. Newton & Peck (2006) listed it (as C. 399; Herman, 2001b: 3326.
maxillosus) as a predator. Other biology and life-
history characteristics are unknown. Pupae and
larvae are unknown. Type material: Staphylinus flavipennis Hope.
Lectotype (here designated). 씸, ‘[circular with red
Etymology: The species epithet ‘galapagensis’ is border] Type| H.T./ flavipennis Hope./ Hardwicke|
derived from a combination of Galapagos and the Bequest./ flavipennis. Hope.| 4241/ FMNH-INS
Latin adjectival suffix -ensis, meaning ‘belonging to’, 0000 016 808/ [red] LECTOTYPE| Staphylinus| fla-
in reference to the species being endemic to the vipennis Hope, 1831| designated by| D. J. Clarke
Galápagos Islands. 2008’ (in BMNH). Only one specimen was in Hard-
wicke’s collection.
Creophilus villipennis Kraatz. Lectotype (here
Remarks: Known literature citations for Creophilus designated). 씹, ‘1307/ [yellow] India sept.| Bacon
from the Galápagos Islands are most likely all C. leg./ Coll. Kraatz/ [red] Syntypus/ coll. DEI|
galapagensis, recorded as C. maxillosus or C. villosus. Eberswalde/ coll. DEI| Müncheburg/ FMNH-INS
The first records are three specimens (not examined 0000 016 810/ [red] LECTOTYPE| Creophilus| vil-
by me) collected by Charles Darwin on Chatham lipennis Kraatz, 1859| designated by| D. J. Clarke
Island (= San Cristóbal) during his famous ‘Beagle’ 2008’ (in DEI). Specimen and terminalia mounted
Expedition, and first reported as ‘Creophilus, nov. on card; left antennomeres 7–11 and protarsal claw
spec.?’ with a description matching mine (Water- missing; right elytron with pinhole close to scutel-
house, 1845). Waterhouse (1877), Van Dyke (1953), lum. Paralectotypes (3). All with labels ‘[red]
and Franz (1985) listed it as C. villosus. Newton & Syntypus/ Coll. Kraatz/ coll. DEI| Eberswalde/ coll.
Peck (2006) listed it as C. maxillosus, noting it may be DEI| Müncheburg/ [yellow] PARALECTOTYPE|
a new species (on advice of A. Smetana). Waterhouse Creophilus| villipennis Kraatz, 1859| designated
(1845) and Mutchler (1925) compared Darwin’s speci- by| D. J. Clarke 2008’: 씹, ‘[yellow] India sept.|
mens to C. maxillosus and C. villosus. Linell (1898) Bacon leg./ Creophilus| villipennis| Kr./ FMNH-INS
listed it as ‘Creophilus species’, noting it is probably 0000 016 809’ (in DEI); 씸, same collecting data and
C. villosus introduced from North America. Van Dyke ‘FMNH-INS 0000 016 811’ (in DEI); 씸, ‘Decan/
(1953: 38) referred to three of the CAS paratypes Saunders/ Creophilus| villipennis Kr./ FMNH-INS
listed above. Franz (1985) recorded nine specimens 0000 016 812’ (in DEI).
(not examined here) from the same localities on Isla Emus violaceus Fauvel. Holotype (by monotypy). 씹,
Isabela as some paratypes. ‘Molucche| Buru X| Beccari 1875/ Typus/ violaceus|
Fauvel/ Emus| violaceus FvL.| 씹/ Museo Civico| di
Genova/ FMNH-INS 0000 016 807/ [red] HOLO-
4. CREOPHILUS FLAVIPENNIS (HOPE)
TYPE| Emus| violaceus Fauvel, 1878| teste D. J.
COMB. NOV., STAT. NOV.
Clarke 2008’ (in MCSN). Fauvel’s (1878a) original
(FIGS 1I, 4E, 10A, 21, 22) description was based on ‘Un seul 씹’.
Staphylinus flavipennis Hope, 1831: 32. Type locality: Emus insularis Fauvel. Lectotype (here desig-
Nepal; Herman, 2001b: 3502 (as nomen dubium). nated). 씹, ‘Batchian/ V. insularis| FvL./ [red text in
Creophilus villipennis Kraatz, 1859: 67. Type local- black box] Ex-Typis/ R.I.Sc.N.B. 17.479| Coll. et det.
ity: ‘India boreali’; Fauvel, 1875: 54; Bugnion, 1911: A. Fauvel/ FMNH-INS 0000 016 805/ [red] LECTO-
135–152, figs 1–10; Bernhauer & Schubert, 1914: 399; TYPE| Emus| insularis Fauvel, 1879| designated
Cameron, 1932: 261; 1933d: 392; Scheerpeltz, 1933: by| D. J. Clarke 2008’ (in IRSNB). Specimen with
1413; Cerruti, 1951: 17; Scheerpeltz, 1976: 92; right mandible extended. Paralectotype. 씸, ‘Ceram/
Smetana & Davies, 2000: 38; Herman, 2001b: 3325; Coll. et det. A. Fauvel| Creophilus| insularis| Fauv.|
Smetana, 2004: 672; new synonymy. R. I.Sc.N.B. 17.479/ [red text in black box] Ex-Typis/
Emus violaceus Fauvel, 1878a: 248. Type locality: FMNH-INS 0000 016 804/ [yellow] PARALECTO-
‘Moluques, Bouro’; 1879: 94; new synonymy. TYPE| Emus| insularis Fauvel, 1879| designated
Emus villipennis; Fauvel, 1878a: 248; Fauvel, 1879: by| D. J. Clarke 2008’ (in IRSNB); specimen missing
94 (as E. villipenni). right middle leg.
Figure 21. Creophilus flavipennis, details of morphology (setae of head, mandibles, antenna, and pronotum omitted). A,
head, dorsal (small male); B, mandibles, dorsal (bases omitted); C, right antenna, dorsal (only apical setae shown); D,
paramere, dorsal; E, apical part of median lobe with fully everted internal sac, right lateral; F, aedeagus with details of
internal sac, right lateral (parameral setae omitted); G, pronotum (small male); H, female internal genitalia, ventral –
arrow indicates subsclerotized or pigmented regions; I, ventral sclerite of internal sac, ventral (lower) and apical (upper);
J, median lobe apex, ventral – arrow indicates internal tooth flanking the groove of the median orifice. Abbreviations: T1,
T2, T3, mandibular teeth. Additional abbreviations as in Figure 3 (male) and Figure 5 (female). Scale bars: I = 0.1 mm;
rest = 1 mm.
Other material examined: 241 specimens. See sup- Description: Measurements (N = 10씹, 10씸). Forebody
porting information, Appendix S1. length: 씹 6.1–10.3 mm, 씸 6.7–8.9 mm. See support-
ing Table S4 for comparison of ranges of male and
female ratios. Head. Head (Fig. 21A) transverse, only
Diagnosis: With characters of the maxillosus-group; slightly widened posteriorly, suborbicular in smaller
eyes occupying most of sides of head (e.g. Fig. 21A); specimens; HW/HL = 1.25–1.53; basal margins
elytra variably yellowish and/or brownish-black; densely setose; dorsal punctuation fine, moderately
vestiture of head, pronotum, most of elytra, legs, dense; eyes very large (EYL/HL = 0.59–0.81), protrud-
apicolateral regions of tergites IV–V, apical fringes ing, occupying sides of head, lateral margins of head
of tergites III–VI and most of ventral side obscured by eye in dorsal view, HL1/HL2 large, much
whitish-grey to golden-brown; tergal chaetotaxic greater in females than in males (씹 = 1.92–8.00,
formula = 6-6-6-6-4-6. 씸 = 7.67–12.00); antennae as in Figure 21C, moder-
774 D. J. CLARKE
ately clavate; antennomeres 1–6 entirely black, 7–11 middle of the elytra, gradually fading yellowish pos-
greyish-black, 11 frequently reddish-brown; apex of teriorly. These specimens have a distinctly greater
antennomere 11 slightly to moderately emarginate contrast between integument and vestiture coloration
medially, each pair of apical setae widely separated, and match the type specimens of C. insularis
one on each side of apex; mandibles as in Figure 21B, (Fauvel). Indian and South East Asian specimens
longer than head in large males, shorter in have distinctly lighter (almost uniformly yellowish)
females (ML/HL 씹 = 0.89–1.47, 씸 = 0.75–0.96), right elytra and elytral vestiture.
mandibular teeth subequal in size, T1 reduced in
smaller specimens. Thorax and abdomen. Pronotum Comparison: Creophilus flavipennis can generally be
(Fig. 21G) slightly transverse (PW/PL = 1.07–1.22), distinguished from other species of the C. maxillosus-
moderately to strongly narrowed posteriorly; PL 1.12– group by the enormous eyes alone (Figs 1I, 21A), but
1.35 ¥ ESL; basolateral margins very shallowly emar- the vestiture patterning is also diagnostic. The
ginate in front of indistinct hind angles; with dense unique male and female genitalia are of parti-
and long whitish peripheral setae and coarse long cular interest for diagnosing this species, as these
whitish vestiture on anterolateral declivities; scutel- characters are constant throughout the range of the
lum with yellowish-brown vestiture; elytra uniformly species.
blackish-brown to almost yellowish, or blackish-
brown anteriorly fading to yellowish-brown posteri- Distribution (Fig. 22): Oriental, widespread: Nepal,
orly, often with faint violet sheen; vestiture dense, India, Sri Lanka, Myanmar, China (one specimen),
evenly golden-yellow or darker with indistinctly Laos, Thailand, Vietnam, Malaysia, Indonesia, Phil-
lighter setae at middle (as in Fig. 1I); wings fully ippines, and east to Tanimbar Islands (Indonesia).
developed, clear yellowish-brown, without black spot Smetana (2004) records it additionally from Kashmir
in medial field between MP3 and MP4 veins; in India. Herman (2001b) records C. insularis (= C.
pterothorax and legs covered with golden or whitish flavipennis) from New Guinea (evidently based on
vestiture; dorsal abdominal vestiture mostly black, Scheerpeltz, 1933), but I could not confirm that it
with golden brown or whitish maculation restricted to occurs there.
apicolateral regions of tergites (and parasclerites)
IV–V, and apical fringes of tergites III–VI; vestiture of Biology and ecology: Many specimens have been
sternites III–VI mostly golden-brown, VII and VIII taken at light (UV, mercury vapour), others from
mostly black in females, in males each with medial Malaise traps and various carrion. Habitat: lowland
patch of golden vestiture; abdominal tergite VII with rainforest to alpine habitats. Altitude: 61–4500 m.
well-developed palisade fringe. Male genitalia. Aedea- Phenology: throughout the year. Other biology and
gus as in Figure 21F; median lobe apex elongate, life-history characteristics are unknown. Larvae and
deeply incised proximal to lateral recurved teeth, the pupae are unknown.
latter blunt, reduced (Fig. 21F, lit); with paired api-
colateral sclerites (as) articulated to internal edge of Remarks: Staphylinus flavipennis was listed by
median orifice, not fused to it; median orifice inter- Herman (2001b: 3502) as a nomen dubium. Hope
nally grooved on dorsal side and with unique internal (1831) provided a valid description: ‘Ater, capite tho-
teeth (Fig. 21J, arrow). Paramere as in Figure 21D. raceque nigro, elytrisque villoso aureis abdomineque
Internal sac inverted as in Figure 21F, everted as in nigro. Long. lin. 7 ½; lat. 2.’, agreeing both with the
Figure 21E; ventromedian spiculose strip (vr) much lectotype of S. flavipennis and the type series of C.
shorter than ventral sclerite (vs), the latter partly villipennis. The name S. flavipennis is therefore a
divided at middle (Fig. 21I). Female internal genita- senior subjective synonym and should be adopted
lia. Internal female genitalia as in Figure 21H; as the valid name because (1) under Article 23.1
vaginal plate (vp) with median sclerotized strip (ss), it has priority over C. villipennis, and (2) the name
produced posteriorly into unique apically bifid C. villipennis, as a junior synonym, does not
process; vaginal fold with two large pigmented meet the usage requirement of Article 23.9.1.2 (ICZN,
regions dorsal to vaginal plate (Fig. 21H, arrow). Cha- 1999).
etotaxy. Elytral discal series with 6–8 macrosetae; Fauvel (1878a) described the large male holotype
procoxa with two macrosetae at distal margin; tergal of Emus violaceus as follows: ‘denticulo laterali
chaetotaxic formula = 6-6-6-6-4-6, inner lateral mac- postico angulum ipsum formante’, referring to the
rosetae absent from tergite VII. basolateral pronotal denticle typical of large males of
the C. maxillosus-group, and ‘capite inter antennas
Variation: Philippine and eastern Indonesian speci- anguste profundeque bifossulato’, referring to two
mens are overall pitchy black, distinctly darker, with accentuated anterior tentorial pits between the
almost black elytra and whitish-grey vestiture at the antennal fossae – an occasional aberration seen in
Figure 22. Distribution of all georeferenced specimens of C. flavipennis (circles).
other species. In describing Emus insularis, Fauvel 5. CREOPHILUS VARIEGATUS MANNERHEIM

(1879) primarily relied on allometric characters of (FIGS 1J, 2C, D, 4A, 23, 24)
both sexes: ‘capite multo minore, in utroque sexu
Creophilus variegatus Mannerheim, 1830: 20. Type
simili . . . mandibulis duplo brevioribus . . . linea
locality: ‘Brasilia ad Rio Janeiro’; Mannerheim, 1831:
basale collo profunde transversim impressa . . . tho-
434; Nordmann, 1837: 21; Laporte de Castelnau,
race . . . haud perspicue denticulato’. All are typical
1840: 173; Gemminger & Harold, 1868: 575; Fauvel,
of small C. flavipennis specimens. In all other
1875: 54; Lynch Arribalzaga, 1884: 129; Lüderwaldt,
respects C. insularis and C. violaceus type specimens
1911: 417; 419; Bernhauer & Schubert, 1914: 399;
are typical of C. flavipennis.
Scheerpeltz, 1933: 1413; Herman, 2001b: 3325.
Five specimens from the Philippines were identified
Staphylinus variegatus; Dejean, 1821: 21 (nomen
as ‘Creophilus semipubescens’. This is presumably a
nudum: Article 12.1); Erichson, 1839: 351; Blan-
Bernhauer manuscript name as I have seen no pub-
chard, 1842: 76; Murray, 1870: 57. 77.
lished reference. One specimen in NMW (FMNH-INS
Emus variegatus; Fauvel, 1877: xxvii.
0000 106 557) bears a ‘cotypus’ label, another in
ZMHB (FMNH-INS 0000 016 806) bears a ‘paratypus’
label. Three specimens in FMNH were identified Type material: Creophilus variegatus Mannerheim.
by Bernhauer as C. semipubescens (FMNH-INS Lectotype (here designated). 씸, ‘씸/ Silberm/
0000 016 564–565, 016 791). The first of these speci- Brasilia/ [red] LECTOTYPE| Creophilus| variega-
mens bears a label in Bernhauer’s handwriting ‘Creo- tus| Mannh.| designated by| A. Newton 1978/ Mus.
philus semipubescens Brh Typ.’. Zool. H : fors| Spec. typ. no. 15232| Creophilus| var-
776 D. J. CLARKE
Figure 23. Creophilus variegatus, details of morphology (setae of mandibles, antenna, pronotum, and abdominal
structures omitted. A, mandibles, dorsal (bases omitted); B, right antenna, dorsal (only apical setae shown; anterior is to
the left); C, paramere, dorsal; D, ventral sclerite of internal sac, ventral (lower) and apical (upper); E, apical part of
median lobe with fully everted internal sac, right lateral; F, aedeagus with details of inverted internal sac, right lateral
(parameral setae omitted); G, right elytron (only whitish setae shown); H, pronotum (small male); I, female internal
genitalia, ventral; J, ventral sclerite of internal sac with attached sclerotized bar (bar), ventral; K, male abdominal
sternites VII and VIII; L, female abdominal tergite X; M, median lobe apex, ventral. Abbreviations: T1, T2, T3,
mandibular teeth. Additional abbreviations as in Figure 3 (male) and Figure 5 (female). Scale bars: D and J = 0.1 mm;
rest = 1 mm.
iegatus Mannh./ FMNH-INS 0000 016 813/ [red] Diagnosis: With characters of the maxillosus-group;
LECTOTYPE| Creophilus| variegatus Mannerheim, antennae subincrassate; apex of antennomere 11
1830| designated by| D. J. Clarke 2008’ (in MZHF). convex (Fig. 23B); pronotum strongly constricted
Newton’s lectotype designation has not been pub- basally with sharply delimited hind angles (Fig. 23H);
lished (A. F. Newton, pers. comm.). elytral epipleura yellowish; with extensive pattern of
white and golden brown vestiture on pronotum,
Other material examined: 333 specimens. See sup- elytra, and abdomen (Fig. 1J); tergal chaetotaxic
porting information, Appendix S1. formula 6-6-6-4(6)-6-8.
Description: Measurements (N = 10씹, 10씸). Forebody Thorax and abdomen. Pronotum (Fig. 23H) slightly
length: 씹 6.4–9.9 mm, 씸 6.6–8.3 mm. See supporting transverse (PW/PL = 1.07–1.22), with sides subparal-
Table S4 for comparison of ranges of male and female lel to slightly convex anteriorly, distinctly constricted
ratios. Head. Head slightly wider anteriorly in large posteriorly, and hind angles sharply delimited; PL
males, subtrapezoidal (wider posteriorly) to subor- 1.23–1.50 ¥ ESL; vestiture of anterolateral declivities
bicular in females and smaller males; HW/HL = 1.25– and sides sparse, whitish-grey and brown; vestiture of
1.56; basal margins setose; dorsal punctation coarse, basal margin golden-brown, dense, and 2¥ as long as
moderately dense; eyes large (EYL/HL = 0.48–0.70), setae on anterior angles, with central setae radiating
dorsolateral in large males, more lateral in females outwards; basolateral impressions broad and deeply
and smaller males, lateral margins of head visible in set, with dense golden-brown setae; vestiture of
dorsal view only in large males, obscured by eye in scutellum whitish-grey anteriorly, brown posteriorly;
smaller specimens, HL1/HL2 greater in females than disc of elytra brown-black, epipleura yellowish; vesti-
males (씹 = 1.54–3.00, 씸 = 3.20–5.00); antennae char- ture of elytral surface dense, with patches of whitish-
acteristic (Fig. 23B), subincrassate; antennomeres grey, brown, and golden setae forming complex
1–3 brownish-black, 4–11 variably yellowish-brown pattern, posterior-most setae forming loose golden
and black; antennomeres 9 and 10 narrowed anteri- apical band (Figs 1J, 23G shows white setae only);
orly, thickened posteriorly; apex of antennomere 11 apical setae 2¥ length of discal setae; hind wings fully
convex medially; each pair of apical setae moderately developed, clear yellowish-brown, without black spot
widely separated, one on each side of apex; mandibles in medial field between MP3 and MP4 veins;
as in Figures 2C, D and 23A, longer than head in pterothorax and legs covered with whitish and golden
large males, shorter in females (ML/HL 씹 = 0.95– vestiture; dorsal abdominal vestiture arranged into
1.48, 씸 = 0.89–0.96), T3 larger than T1 and T2. characteristic pattern of whitish-grey, golden, and
Figure 24. Distribution of all georeferenced Creophilus specimens from South America. C. variegatus (triangles),
C. erythrocephalus (squares), C. maxillosus maxillosus (circles).
778 D. J. CLARKE
Figure 25. Creophilus erythrocephalus, details of morphology (setae of mandibles, antenna, and pronotum omitted).
A, pronotum (large male); B, mandibles, dorsal (bases omitted); C, right antenna, dorsal (only apical setae shown);
D, paramere, dorsal. Abbreviations: T2, T3, mandibular teeth. Scale bars = 1 mm.
brown maculation (Fig. 1J); vestiture on all sternites Distribution (Fig. 24, triangles): Endemic to South
mostly whitish, with irregular black patches laterally; America: Argentina, Bolivia, Brazil, Chile (one
abdominal tergite VII with well-developed palisade record), Paraguay, Peru (one record), Uruguay.
fringe. Male genitalia and secondary sexual charac-
ters. Sternite VII distinctly emarginate medioapically Biology and ecology: Lüderwaldt (1911) records the
(Fig. 23K). Aedeagus as in Figure 23F; median lobe species from carrion, and Lynch Arribalzaga (1884)
apex long, pointed (Fig. 23M); paired apicolateral additionally reports predation of calliphorid maggots.
sclerites (as) fused to sides of median lobe. Paramere Habitat: unknown. Altitude: sea-level to 3000 m.
as in Figure 23C; with apex entire; and with setae Phenology: throughout the year. Other biology and
along entire length. Internal sac inverted as in life-history characteristics are unknown. Larvae and
Figure 23F, everted as in Figure 23E, paired basoven- pupae are unknown.
tral spiculose regions absent; with additional sclero-
tized bar (Fig. 23J) connecting ventral sclerite (vs)
Remarks: Dejean (1821) listed Staphylinus variegatus
with base of copulatory piece (cp); copulatory piece
Dej., attributing the name to himself, but did not
sclerites mostly fused along midline, without mem-
provide a description. Mannerheim (1830) published
branous sheath (Fig. 4A). Female internal genitalia.
the name Creophilus variegatus Dej., attributing the
Apex of tergite X narrowly rounded (Fig. 23L). Inter-
name to Dejean, but also provided the first description
nal female genitalia as in Figure 23I; vaginal plate
so authorship is attributed to him. Mannerheim (1830)
(vp) rounded posteriorly, without median sclerotized
did not indicate the number of specimens examined.
strip. Chaetotaxy. Basiantennal and basisternal mac-
rosetae absent; elytral discal series with 4–6 macrose-
tae; mesocoxae with one macroseta at distal margin; CREOPHILUS ERYTHROCEPHALUS SPECIES-GROUP
tergal chaetotaxic formula 6-6-6–4(6)-6-8, inner later- This species group was originally proposed by Newton
als present or absent on tergite VI. (1985) for the austral species. With removal of one
species from synonymy and description of one new
Comparison: Creophilus variegatus is easily distin- species, seven species are included here.
guished from other species of the C. maxillosus-group
by the body vestiture patterning (Fig. 1J), antennae Diagnosis: Head at least partly orange-red or
(Fig. 23B), and sharply delimited basal pronotal yellowish-brown (Fig. 1A–E), dorsally with black spot
angles (Fig. 23H). Males are unusual within the variable in size and shape (Fig. 7); antennomere 2
genus in having the apex of sternite VII distinctly subequal in length to 3 or longer; antennomere 11
emarginate medially (Fig. 23K) and the parameral with apical setae close together anterior to antennal
apex entire (Fig. 23C). axis (e.g. Fig. 25C); right mandible with T3 proximal
to T2 and base of mandibular blade, mostly to fully 1972a, b, 1974; Bellas, Brown & Moore, 1974; Coiffait,
visible dorsally (e.g. Fig. 25B); anterior pronotal 1976: 218; Legner, 1978: 348, 352; Radford, 1981: 174;
angles converging (Fig. 25A); abdominal tergites Frank, 1982: 34; Matthews, 1982: 7, fig. 34 (cover
III–V with secondary transverse basal ridge (Fig. 2L, picture); Andrews & Gibbs, 1989: 107, 113; Sáiz,
sbr); marginal fringe of tergites III–VI and body ves- Solervicens & Ojeda, 1990: 50, fig. 16 (as erytroceph-
titure uniformly black (or golden-brown in most C. alus); Kukalová-Peck & Lawrence, 1993: 236, fig. 46;
huttoni specimens); median lobe of aedeagus continu- Naumann, 1993: 163; (common name as devil’s coach-
ously sclerotized ventrally (Fig. 3A). Large males with horse) Lawrence & Britton, 1994: 11, 97, fig. 5f, pl.
apical portion of mandibles less than or equal to 0.9 10c; Nishida, 1994: 71; Desender & Baert, 1996: 43;
times length of basal portion (ML1/ML2 range of all Newton, 1997: 25; Smithers, 1998: 16; Herman,
males = 0.58–0.90); with ventrolateral carina of head 2001b: 3316; Levot, 2003: 32; Archer, 2004; 35;
(Figs 2A, 31B, vl); pronotum widest slightly anterior Nowak, 2004: 13; Read & Wilson, 2004: 54; Hawkes-
to middle (e.g. Fig. 25A); base of profemur without wood & Turner, 2008.
spine-like apophysis; superior pronotal marginal line
not deflected beneath pronotum basally, basolateral Type material: Staphylinus erythrocephalus Fabri-
denticle absent (Fig. 25A). cius. Lectotype (designated in footnote by Steel,
1949: 58, labelled by me). 씸, ‘Staph. erythroceph.|
Fab. Entom.p.265.6/ Creophilus| erythrocephalus|
6. CREOPHILUS ERYTHROCEPHALUS (FABRICIUS)
F.| det. A.F. Newton 1989/ FMNH-INS 0000 016 779/
(FIGS 1A, 2A, B, L, 3A, B, G, H, O, 4H, [red] LECTOTYPE| Staphylinus| erythrocephalus
5B, 7A, 10C, 24, 25, 26, 28) Fabricius, 1775| des. by W.O. Steel, 1949, teste | D.J.
Staphylinus erythrocephalus Fabricius, 1775: 265. Clarke 2008’ (in Banks Collection, BMNH). Specimen
Type locality: ‘nova Hollandia’; Goeze, 1777: 724; Fab- glued to pin ventrally, wings unfolded, missing left
ricius, 1781: 335 (not seen); Fabricius, 1787: 220; antennomeres 3–11, right prothoracic leg missing
Fabricius, 1793: 523; Olivier, 1795: (42): 12, pl.2, tibia and tarsus, but coxa and femur attached to
fig. 9; Fabricius, 1801: 593; Turton, 1802: 511; specimen with glue. Paralectotypes (2). 1씸, no data
Latreille, 1804: 293; Gravenhorst, 1806: 127; Dejean, but with labels ‘Creophilus| lanio Er.| det. A.F.
1821: 21; Boisduval, 1835: 55; Erichson, 1839: 351; Newton 1989/ FMNH-INS 0000 016 780/ [yellow]
Blanchard, 1842: 77; Fairmaire, 1849: 288; Murray, PARALECTOTYPE| Staphylinus| erythrocephalus
1870: 57, 89; Steel, 1949: 57, 58 (synonym of C. lanio Fabricius, 1775| des. by W.O. Steel, 1949, teste | D.J.
‘ex parte’); Zimsen, 1964: 231; Radford, 1981: 174 Clarke 2008/ Creophilus lanio (Erichson, 1839) det.
(invalid holotype designation). D.J. Clarke 2006’ (in Banks Collection, BMNH).
Staphylinus unipunctatus Hope, 1831: 32. Type Specimen pinned through base of abdomen, glued to
locality: Nepal; Lea, 1925: 229 (synonym of C. eryth- pin ventrally, missing right metathoracic leg, and
rocephalus); Steel, 1949: 57 (synonym of C. erythro- with wings unfolded but partly disintegrated. 1씹, no
cephalus); Radford, 1981: 174 (as C. punctatus, error data but with labels ‘[yellow] PARALECTOTYPE|
for unipunctatus); Herman, 2001b: 3316 (synonym of Staphylinus| erythrocephalus Fabricius, 1775| des.
C. erythrocephalus). by W.O. Steel, 1949, teste | D.J. Clarke 2010/ Creo-
Emus erythrocephalus; Dejean, 1833: 59 (not seen); philus lanio (Erichson, 1839) det. D.J. Clarke 2010’
Fauvel, 1877: 249; Fauvel, 1878a: 248; Fauvel, 1878b: (in ZMUC). Specimen pinned in unit tray with label
541; Fauvel, 1889: 261. pinned in tray: ‘erythro| cephalus’ [Fabricius’s hand-
Creophilus erythrocephalus; Nordmann, 1837: 23; writing]. The lectotype and female paralectotype are
Motschulsky, 1858a: 49; Fauvel, 1875: 56; Olliff, 1887: pinned with thick pins typical of the period (Andrews
492; Olliff, 1889: 78, 81; Fauvel, 1903: 267; Froggatt, & Gibbs, 1989). The male paralectotype (ZMUC) has
1907: 137; Bernhauer, 1908: 19; Sharp & Muir, 1912: a hole in the left elytron consistent with the size of
498; Bernhauer & Schubert, 1914: 398; Bernhauer, the lectotype pin, but was at some point re-pinned
1920: 16; Fullaway, 1923: 185; Swezey, 1923: 303; with a modern pin through the right elytron. Steel’s
Cameron, 1924: 87 (probable error for C. imitator; (1949) lectotype designation is valid under Article
specimen not seen); Lea, 1925: 229; Tillyard, 1926: 74.5 (ICZN, 1999). Both paralectotypes are specimens
209; Cameron, 1933: 83; Fuller, 1934: 14, 18; of C. lanio.
Cameron, 1937: 104 (error for C. imitator; see Distri- Staphylinus unipunctatus Hope. Lectotype (here
bution section, below); Hawkins, 1942: 885; Curran, designated). 씸, ‘[circular with red border] Type|
1945: 174; MacKeown, 1944: 127; Steel, 1949: 57, 59, H.T./ unipunctatus. Hope/ Hardwicke| Bequest./
60, 61, figs 1, 4, and 9; Gourlay, 1950: 186; Bornemis- unipunctatus. Hope.| 4242 [in pencil] erythroceph-
sza, 1957: 6; Coiffait & Sáiz, 1968: 364, figs 3 and 4; alus Fabr. Hopes locality wrong/ FMNH-INS
Sáiz, 1971: 349 (as erytrocephalus); Meyer-Rochow, 0000 016 783/ [red] LECTOTYPE| Staphylinus| uni-
780 D. J. CLARKE
punctatus Hope, 1831| designated by| D.J. Clarke Paramere as in Figure 25D. Internal sac inverted as in
2008’ (in BMNH). Hope did not mention the number Figure 3A and B, everted as in Figure 3H; paired
of specimens examined. Although only a single speci- apicolateral sclerites (as) separated from sclerotized
men was in Hardwicke’s collection, there is insuffi- median lobe by distinct membranous strip; ventral
cient evidence to consider it the holotype, as labelled. sclerite of internal sac flattened (Fig. 3G), without
conspicuous sculpture. Female internal genitalia.
Other material examined: 1513 specimens. See sup- Internal female genitalia as in Figure 5B; vaginal
porting information, Appendix S1. plate with paired lateral sclerites (pls), posterolateral
aspects membranous; vaginal fold (vf) with pair of
Diagnosis: With characters of the erythrocephalus- separated sclerites dorsal to posterolateral lobes of
group; head orange-red, with transverse diameter of vaginal plate. Chaetotaxy. Elytral discal series with
circular medial black spot greater than 0.5 ¥ EYL three macrosetae; metaventrital macroseta present;
(Figs 1A, 7A); right mandible with two distinct teeth tergal chaetotaxic formula = 4-6-6-6(4)-4-6, medial
(Fig. 25B); elytra metallic blue, humeri glabrous, pair of macrosetae absent on tergite III, inner lateral
shining black; abdomen uniformly black; tergal cha- macrosetae present or absent on tergite VI, absent on
etotaxic formula = 4-6-6–6(4)-4-6. tergite VII.
Description: Measurements (N = 10씹, 10씸). Forebody Comparison: Creophilus erythrocephalus is most

length: 씹 5.1–9.4 mm, 씸 5.3–7.8 mm. See supporting similar to C. imitator and C. lanio, but the metallic
Table S5 for comparison of ranges of male and female blue elytra with shining black humeri and large,
ratios. Head. Head (Figs 2A, B, 7A) orange/red, nar- distinctly circular black cranial spot are diagnostic.
rowly black around mouthparts and antennal fossae, In addition, C. erythrocephalus may be easily distin-
vertex with large circular and sharply defined black guished from C. lanio by its black ninth abdominal
spot, situated posterior to and not concealing dorsal segment.
tentorial pits, with transverse diameter greater than
0.5¥ EYL; strongly trapezoidal, much wider posteri- Distribution (Figs 24, 26, 28): Herman (2001b)
orly; HW/HL = 1.37–1.69; shining, without distinct recorded C. erythrocephalus from Nepal, New Guinea,
microsculpture; eyes small to moderately large (EYL/ Australia, Lord Howe Island, New Caledonia, Tonga,
HL = 0.37–0.53), dorsolateral; lateral margins of head Society Islands (French Polynesia), Hawaii, Easter
visible in dorsal view (not obscured by eye); HL1/HL2 Island, and Chile. It is apparently native to Australia
greater in females than males (씹 = 1.38–1.83, and introduced to Chile (Fauvel, 1903) where it is
씸 = 1.50–2.20); antennae as in Figure 25C, antenno- well established [earliest records from 1898 (Sáiz,
meres 1–6 black, 7–11 greyish-black, 11 as long as or 1971), in MNNC]. It was deliberately introduced to
very slightly longer than 9–10 together; mandibles as Hawaii (Fullaway, 1923; Swezey, 1923) but never
in Figure 25B, moderately longer than head in large established there (Newton, 1997). Old New Zealand
males, subequal to head in females (ML/HL 씹 = 0.84– records are doubtful (e.g. Fairmaire, 1849; Murray,
1.24, 씸 = 0.81–1.00), right mandible with two teeth, T1 1870; Curran, 1945; three old specimens labelled
absent, T3 much larger than T2; left mandible with ‘Neuseeland’) and it is certainly absent from there at
ventral basolateral ridge only weakly developed. present. The Nepal locality for the S. unipunctatus
Thorax and abdomen. Pronotum (Fig. 25A) slightly lectotype is almost certainly incorrect (see label data
transverse (PW/PL = 1.10–1.19); PL 1.24–1.57 ¥ ESL; quoted above); no other Nepalese records were seen.
with basolateral margins very slightly emarginate, It also occurs in Fiji (five specimens, Taveuni Island).
hind angles indistinct; with short and sparse periph- Historical New Guinean records are probably misla-
eral vestiture, and dense longer vestiture on antero- belled: a single specimen (‘1 example. Doormanpad-
lateral declivities; elytra, except humeri, metallic blue bivak 1410 m. Oct. 1920’) collected by W. C. van
or violet, densely setose, and only very slightly rugose; Heurn during his 1920–21 expedition to Dutch North
humeral regions black, shining, distinctly callused; New Guinea was recorded as C. erythrocephalus by
wings fully developed, black with distinct black spot in Cameron (1924) (probably in Zoological Museum,
medial field between MP3 and MP4 veins; abdomen Amsterdam). Later, Cameron (1937) listed another
shining and uniformly black; tergite VII with well- New Guinean specimen (Mt Tafa, alt. 8500 ft., ii.1934;
developed palisade fringe. Male genitalia and second- FMNH-INS 0000 016 773, in BMNH) that I deter-
ary sexual characters. Sternite VII with impressed mined to be C. imitator. I have seen one recent
transverse region of increased setal density, containing specimen from New Guinea (‘Lae, TPNG, VIII 59,
high density of micropunctures. Aedeagus as in A.C. Robinson, UQIC Reg.#80897) and there is
Figure 3A, B; median lobe apex produced distinctly nothing to suggest it is mislabelled (TPNG = Territory
dorsad into short and broad globular process (Fig. 3O). of Papua New Guinea).
Figure 26. Distribution of all georeferenced Australian specimens of C. erythrocephalus (circles). See Figure 24 for
distribution in South America, and Figure 28 for New Guinea. This species is also known from Lord Howe Island, Fiji,
French Polynesia, New Caledonia, Tonga, and Vanuatu.
Biology and ecology: Creophilus erythrocephalus Remarks: Fabricius did not indicate the number of
prefers open or disturbed habitats, and typically is specimens examined, or designate a ‘type’. Zimsen
found in synanthropic situations. Many specimens (1964: 231) listed two type specimens for Staphylinus
have been taken at light (UV, blacklight), dung, and erythrocephalus, one in Kiel, one in London (BMNH:
carrion of various sorts. Habitat: Banksia and Banks Collection). The former specimen was trans-
Eucalyptus forest/scrub; sclerophyll forest and wood- ferred to ZMUC, along with the rest of Fabricius’
land, coastal sand heath, sand dunes, and beach. collection, and the existence of a type in his personal
Altitude: sea level to 1600 m. Phenology: throughout collection is reasonable as ‘Banks was a generous
the year. Larvae and pupae are known and will be man . . . Fabricius certainly added specimens from
described elsewhere. Creophilus erythrocephalus is a the collections he studied to his own collection’
known predator of fly larvae, and has been tested as (Radford, 1981: 157). C. O. Waterhouse studied the
a biocontrol agent for flies in Hawaii (Fullaway, Banks Collection (BMNH) in the late 19th century,
1923; Swezey, 1923). Dynamics of arrival and and, according to Radford (1981: 156), he applied type
departure at carrion have been recorded in labels to specimens he was reasonably sure were
Australia (Fuller, 1934; Bornemissza, 1957; Levot, those to which Fabricius’s descriptions referred.
2003; Archer, 2004). Results of these studies There are, however, two specimens in the Banks
are consistent with those discussed under C. max- Collection (BMNH), and based on their aforemen-
illosus, above. Fuller (1934) noted that larvae, while tioned details, both of these and the ZMUC specimen
developing, can remain at a carcass for periods can be inferred to have been studied by Fabricius and
exceeding 20 days and that, in agreement with should be considered syntypes. It is possible that
other studies, adults tend to occur at a carcass only Zimsen did not list the second BMNH specimen
intermittently. because Steel (1949) previously identified it as C.
782 D. J. CLARKE
lanio. Radford (1981) also studied the Fabricius type situated on a sparsely overgrown summit, less cloudy
material, but considered the now lectotype specimen [than Mist Camp]’. He provided a map illustrating
to be a holotype, probably basing this on the original the location of Top and Mist Camps in the Snow
description, C. O. Waterhouse’s presumably attached Mountains (p. 272).
‘type’ label, and by exclusion of the C. lanio specimen.
According to Radford (1981: 174), the lectotype should Other material examined: 91 specimens. See support-
have had a second label with ‘BMNH Staphylinus ing information, Appendix S1.
erythrocephalus’, and a museum (Waterhouse’s?) type
label. No such labels were present on either specimen Diagnosis: With characters of the erythrocephalus-
although the type label pinned in the drawer next to group; head orange-red, with transverse diameter of
the specimens could have once been applied to the circular medial black spot less than 0.5¥ EYL
lectotype. (Fig. 7D); right mandible with two distinct teeth
(Fig. 27A); pronotal basolateral impressions present
(Fig. 27E); elytra and abdomen black; basiantennal
7. CREOPHILUS IMITATOR CAMERON and basolateral pronotal macrosetae absent; tergal
(FIGS 4I, 7D, 9A, B, 27, 28) chaetotaxic formula 6-6-6-4-2–4.
Creophilus imitator Cameron, 1952: 255. Type local- Description: Measurements (N = 10씹, 10씸). Forebody
ity: ‘New Guinea (Irian Jaya), Snow Mountains, Top length: 씹 7.0–10.2 mm, 씸 7.1–8.8 mm. See supporting
Camp, 2100 m’; Herman, 2001b: 3317. Table S5 for comparison of ranges of male and female
ratios. Head. Head orange-red, narrowly black around
Type material: Creophilus imitator Cameron. Holo- mouthparts and antennal fossae, with small circular
type (original designation). 씹, ‘Neth. Ind. – Ameri- and sharply delimited black spot situated posterior to
can| New Guinea Exped.| Top Camp 2100 m| and not concealing dorsal tentorial pits, with trans-
25.I.1939 L.J. Toxopeus/ [blue] typus/ [black- verse diameter less than 0.5¥ EYL (Fig. 7D); black and
bordered] MUSEUM LEIDEN| Creophilus| imita- orange coloration strongly contrasting; strongly trap-
tor| Cam.| det. M. Cameron/ M. Cameron det., 19| ezoidal and much wider posteriorly in large males,
Creophilus| imitator| [red ink] TYPE [black ink] almost subrectangular in females and smaller males;
Cam/ [red] Holotype/ [red] HOLOTYPE| Creophilus| HW/HL = 1.43–1.70; surface dull, with distinctive
imitator Cameron, 1952| teste D.J. Clarke 2010’ (in micropunctate microsculpture and very shallow indis-
RMNH). Pinned through right elytron; missing both tinct and moderately dense impressions (Fig. 9A, B);
antennae; wings out, intact; genital segment (in two eyes large, protruding somewhat laterally (EYL/
parts) and aedeagus dissected (by Cameron?) and HL = 0.40–0.55), lateral margins of head only visible in
mounted on old card beneath specimen; pinned with dorsal view in largest males, obscured by eye in
brass-headed pin. Paratypes (4). 1씹, ‘[circular with smaller specimens; HL1/HL2 greater in females than
yellow border] Para-| type/ Neth. Ind. –American| males (씹 = 1.13–1.88, 씸 = 1.71–2.33); antennae as in
New Guinea Exped.| Mist Camp 1800 m| Figure 27B; antennomeres 1–6 black, 7–11 greyish-
18.I.1939 L.J. Toxopeus| op rottend aas [on rotting black, 11 elongate, as long as or longer than 9–10
carrion]/ 씹/ M. Cameron det., 19| Creophilus| imi- together; mandibles as in Figure 27A, moderately
tator| P-TYPE Cam./ M. Cameron.| Bequest.| B.M. longer than head in large males, slightly shorter than
1955–147/ imitator/ FMNH-INS 0000 016 771’; 1씸, head in females (ML/HL 씹 = 0.86–1.22, 씸 = 0.89–
same collecting data, ‘M. Cameron.| Bequest.| B.M. 0.96), right mandible with two teeth, T3 much larger
1955–147/ FMNH-INS 0000 016 772’ (in BMNH); 1씹, than T2. Thorax and abdomen. Pronotum characteris-
aedeagus and genital segment in glycerin, same col- tic (Fig. 27E), subquadrate (PW/PL = 1.03–1.16), in
lecting data, except no collecting method, ‘[blue] females wider at hind angles than anterior angles;
Paratypus/ Creophilus| imitator| Cam.| det. M. sides subparallel, almost straight to only slightly
Cameron/ FMNH-INS 0000 012 322’ (in AMNH); 1씸, convex in large males; PL 1.12–1.33 ¥ ESL; basolateral
‘Neth. Ind.-American| New Guinea Exped.| Top margins very slightly emarginate, hind angles dis-
Camp 2100 m| 30.I.1939 L.J. Toxopeus/ ‘[Cameron’s tinctly rounded; with sparse peripheral setae and
handwriting] Creophilus| imitator Cam.| paratype| short, sparse vestiture on lower anterolateral declivi-
dt. Cameron ′51’ (in MBBJ). ties; shallow basolateral impressions present, smooth
and asetose (Fig. 27E); elytra distinctly widened pos-
teriorly, uniformly black, sparsely and finely setose,
Description of type locality: Top Camp, in the Snow finely rugose, especially around humeral regions;
Mountains (Indonesia: Papua), is described by humeri glabrous, impunctate, not distinctly callused;
Toxopeus (1940: 279) as ‘an outpost of Mist Camp, wings fully developed, clear yellowish-brown, without
Figure 27. Creophilus imitator, details of morphology (setae of mandibles, antenna, and pronotum omitted). A, man-
dibles, dorsal (bases omitted); B, right antenna, dorsal (only apical setae shown); C, median lobe apex, ventral; D,
aedeagus with details of inverted internal sac, right lateral (parameral setae omitted); E, pronotum (female); F, female
internal genitalia, ventral – arrow indicates small sclerotic ridges of vaginal fold; G, paramere, dorsal; H, ventral sclerite
of internal sac, ventral (lower) and apical (upper). Abbreviations: T2, T3, mandibular teeth. Additional abbreviations as
in Figure 3 (male) and Figure 5 (female). Scale bars: H = 0.1 mm; rest = 1 mm.
black spot in medial field between MP3 and MP4 veins; branous strip. Paramere as in Figure 27G, widely
abdomen shining and uniformly black; vestiture fine, emarginate apically. Internal sac inverted as in
moderately dense; microsculpture extremely fine, pro- Figure 27D. Female internal genitalia. Internal female
ducing weak iridescent reflection, particularly on genitalia as in Figure 27F; vaginal plate (vp) without
ventral side; tergite VII with well-developed palisade median sclerotized strip, with paired lateral sclerites
fringe. Male genitalia and secondary sexual characters. (pls); vaginal fold (vf) with pair of separate sclerites
Sternites VI and VII of males with scattered dorsal to posterolateral lobes of vaginal plate, with
micropores, those on sternite VII situated in asetose small sclerotic ridges medially (Fig. 27F, arrow). Cha-
impressed transverse region. Aedeagus as in etotaxy. Basiantennal and basolateral pronotal mac-
Figure 27D; median lobe with apex produced obliquely rosetae absent; elytral discal series with three or four
dorsad into short and narrow globular process macrosetae; tergal chaetotaxic formula 6-6-6-4-2–4,
(Fig. 27C), with paired apicolateral sclerites (as) sepa- tergites VI–VIII without inner lateral macrosetae,
rated from sclerotized median lobe by distinct mem- tergite VII without medial macrosetae.
784 D. J. CLARKE
Figure 28. Distribution of all georeferenced specimens of C. albertisi (circles), C. erythrocephalus (square) and C. imitator
(triangles) in New Guinea.
Comparison: Creophilus imitator has been histori- Remarks: The original description indicates that
cally confused with C. erythrocephalus but is readily Cameron saw five specimens from two localities: ‘Top
distinguished from that species by the much smaller Camp, 2100 m, 25.i.39 (Type); Mist Camp, 1800 m,
black spot on the head and the combination of black 18.i.39, L.J. Toxopeus. Five specimens’. According to
finely rugose elytra and basolateral pronotal impres- the editor of the original description, ‘types’ in Cam-
sions. The distinctive micropunctate microsculpture eron’s paper were deposited in ‘Leiden Museum’
of the head (shared with C. albertisi) also distin- (RMNH), with paratypes and duplicates deposited in
guishes it from both C. erythrocephalus and C. lanio. ‘Museum Zoologicum Bogoriense’ (MBBJ) and ‘in the
author’s collection’ (in BMNH). No details about how
a paratype came to be deposited in AMNH could be
Distribution (Fig. 28, triangles): New Guinea: Papua determined but this is not inexplicable, as Richard
New Guinea, Irian Jaya (Indonesia). Archbold, who funded the 1939 New Guinea expedi-
tion, was affiliated with AMNH.
Last (1987) recorded Creophilus from New Guinea,
Biology and ecology: Rare in collections; most speci-
but did not list the species, nor did he provide any
mens without habitat or collection data. Twenty speci-
records.
mens were collected from dung traps set in montane
rainforest. Other records are from mercury vapour
8. CREOPHILUS LANIO (ERICHSON)
light traps and carrion. Habitat: montane rainforest.
Toxopeus (1940) recorded details of the habitat at (FIGS 2H–J, 3Q, 4J, 7B, 29, 30)
Mist Camp as ‘on the mountain ridge S. of Bernhard Staphylinus lanio Erichson, 1839: 352. Type locality:
Camp on the Idenburg R. Dense, very damp forest ‘Terra Van-Diemenii’; Gravenhorst, 1806: 126.
in a saddle’. Altitude: 1200–2591 m. Phenology: Staphylinus oculatus var. Gravenhorst; Dejean,
January–March, June–December. Other life-history 1821: 21; Erichson, 1839: 352 (synonym of C. lanio).
characteristics are unknown. Larvae and pupae are Creophilus lanio; Fauvel, 1875: 56; Bernhauer &
unknown. Schubert, 1914: 398; Lea, 1925: 229 (synonym of
Figure 29. Creophilus lanio, details of morphology (setae of mandibles, antenna, and pronotum omitted). A, mandibles,
dorsal (bases omitted); B, right antenna, dorsal (only apical setae shown); C, female internal genitalia, ventral; D,
paramere, dorsal; E, aedeagus with details of inverted internal sac, right lateral (parameral setae omitted); F, pronotum
(female); G, ventral sclerite of internal sac, ventral (lower) and apical (upper); H, basal cap of internal sac; I, median lobe
apex, ventral. Abbreviations: T1, T2, T3, mandibular teeth. Additional abbreviations as in Figure 3 (male) and Figure 5
(female). Scale bars: G and H = 0.1 mm; rest = 1 mm.
erythrocephalus); Steel, 1949: 58, figs 2, 5, 7 and 8; [lilac] Hist.-Coll. (Coleoptera)| Nr. 5886| Staphylinus
Radford, 1981: 174 (as C. lania, error for lanio); lanio Erichs.| Terra v. Diem., Schayer| Zool. Mus.
Herman, 2001b: 3317. Berlin/ [red] SYNTYPUS| Staphylinus| lanio Erich-
Emus lanio; Fauvel, 1877: 250; Fauvel, 1878a: 248; son, 1839| labeled by MNHUB 2004/ [yellow] PARA-
Fauvel, 1878b: 541. LECTOTYPE| Staphylinus| lanio Erichson, 1839|
Creophilus erythrocephalus var. lanio; Olliff, 1887: designated by| D.J. Clarke 2008’: 2씹, ‘FMNH-INS
492–493. 0000 016 775’; ‘FMNH-INS 0000 016 777’; 1씸,
‘FMNH-INS 0000 016 776’ (in ZMHB).
Type material: Staphylinus lanio Erichson. Lectotype Other material examined: 2743 specimens. See sup-
(here designated). 씸, ‘[orange] Type/ 5886/ [grey] porting information, Appendix S1.
lanio| Er.| terra van Diem [Tasmania], Sch/ [lilac]
Hist.-Coll. (Coleoptera)| Nr. 5886| Staphylinus lanio Diagnosis: With characters of the erythrocephalus-
Erichs.| Terra v. Diem., Schayer| Zool. Mus. Berlin/ group; head orange-red, with large diffuse subquad-
[red] SYNTYPUS| Staphylinus| lanio Erichson, rate or cordiform black spot (Fig. 7B); right mandible
1839| labeled by MNHUB 2004/ FMNH-INS with three teeth (Fig. 29A); elytra black, humeri gla-
0000 016 774/ [red] LECTOTYPE| Staphylinus| brous, conspicuously sculptured; abdominal segment
lanio Erichson, 1839| designated by| D.J. Clarke IX orangeish-brown; parasutural 1 and humeral
2008’ (in ZMHB). Specimen missing left mesotarsus. macrosetae absent, tergal chaetotaxic formula
Paralectotypes (3). All with labels ‘[orange] Type/ = 4-6-6–6-4-6.
786 D. J. CLARKE
Description: Measurements (N = 10씹, 10씸). Forebody

length: 씹 5.5–9.8 mm, 씸 5.3–7.8 mm. See supporting
Table S5 for comparison of ranges of male and female
ratios. Head. Head orange-red, narrowly black around
mouthparts and antennal fossae, with large diffuse
irregularly subquadrate or cordiform black spot
(Fig. 7B), usually concealing dorsal tentorial pits, fre-
quently with midlongitudinal line of black pigment
continuing to frontoclypeal margin and to neck;
strongly trapezoidal, much wider posteriorly;
HW/HL = 1.35–1.59; shining, without distinct micros-
culpture; eyes small to moderately large (EYL/
HL = 0.38–0.56), dorsolateral, lateral margins of head
visible in dorsal view (not obscured by eye); HL1/HL2
greater in females than males (씹 = 1.21–1.83,
씸 = 1.63–2.00); antennae as in Figure 29B, antenno-
meres 1–6 black, 7–11 greyish-black, 11 as long or
very slightly longer than 9–10 together; mandibles as
in Figure 29A, moderately longer than head in large
males, subequal to head in females (ML/HL 씹 = 0.81–
1.32, 씸 = 0.82–1.05), right mandible with three teeth,
T3 largest. Thorax and abdomen. Pronotum
(Fig. 29F) slightly transverse (PW/PL = 1.09–1.25);
PL 1.22–1.45 ¥ ESL; with basolateral margins dis-
tinctly emarginate, hind angles distinct; with sparse
peripheral setae and short, sparse vestiture on lower
anterolateral declivities; elytra uniformly black,
humeral regions shining, distinctly callused, disc
densely setose, sparsely rugosely sculptured, espe-
cially on humeri; wings fully developed, black with Figure 30. Distribution of all georeferenced specimens of
distinct black spot in medial field between MP3 and C. lanio in south eastern Australia and Tasmania (circles).
MP4 veins; abdomen shining black, except for lightly One possibly mislabeled specimen was seen from Western
pigmented orangeish-brown ninth segment; tergite Australia.
VII with well-developed palisade fringe; styli bico-
loured, orangeish-brown basally and black distally.
Male genitalia and secondary sexual characters. Ven- Variation: The dorsomedial spot in C. lanio is very
trolateral carina of large males indistinct, partially variable. Occasionally, usually in smaller specimens,
obliterated by punctures and secondary anastomosing it is expanded to cover most of the area between the
ridges. Aedeagus as in Figure 29E; median lobe apex eyes, although always surrounded by the usual
extended into long blunt point (Fig. 29I), produced orange coloration of the head.
slightly dorsally at tip, with paired apicolateral
sclerites (as) separated from sclerotized median Comparison: Creophilus lanio may be immediately
lobe by distinct membranous strip. Paramere as in distinguished from C. erythrocephalus and C. imitator
Figure 29D. Internal sac inverted as in Figure 29E; by the diffuse, never circular cranial spot and by the
ventral sclerite (vs) small, wider than long, distinctly lightly pigmented ninth abdominal segment.
notched apically (Fig. 25G). Female internal genitalia. Contrary to Steel (1949: 58), all specimens I examined
Internal female genitalia as in Figure 29C; vaginal had uniformly shining black elytra, none with a
plate with paired lateral sclerites (pls), posterolateral ‘purplish reflection’. Although Cameron (1952: 255)
areas membranous; vaginal fold forming large finely described the elytral sculpturing of C. imitator as
rugose sclerite (Fig. 29C, vf). Chaetotaxy. Elytra ‘less rugose than in lanio’ it is more distinctive in
without parasutural 1 and humeral macrosetae; C. imitator.
elytral discal series with 3–4 macrosetae; metaven-
trital macroseta absent or undetected; tergal chaeto- Distribution (Fig. 30): Australia: New South Wales,
taxic formula = 4-6-6-6-4-6, medial macrosetae absent Tasmania, Victoria, southern Queensland. One record
on tergite III, inner lateral macrosetae absent on from Western Australia (no locality given, possibly
tergite VII; second gonocoxal macroseta absent. mislabelled).
Biology and ecology: Creophilus lanio is known mostly aedeagus on card, ‘씹/ Albertisi| Fauv./[blue] ex coll.|
from forested areas, with few records from pasture Klima/[blue] ex coll.| Scheerpeltz/ Albertisi|
and coastal habitats. Most collections are from Fauv./ FMNH-INS 0000 012 299’ (in NMW). Fauvel’s
carrion-baited pitfall traps. Specimens have also been description was based on numerous specimens col-
taken at light, in yellow pan traps, in Malaise and lected by D’Albertis: ‘Nouvelle-Guinée, Fly River; en
window traps, and in dung of various sorts. Habitat: nombre (L.M. D’Albertis).’ These were deposited in
drier woodlands to temperate rainforests of many ‘Collection de Musee Civique de Genes et la mienne’
kinds (e.g. Acacia, Banksia, Eucalyptus, Nothofagus, (MCSN and Fauvel’s collection).
and Pomaderris). Altitude: sea level to 1650 m. Phe-
nology: throughout the year. Other biology and life- Other material examined: 59 specimens. See support-
history characteristics are unknown. Larvae and ing information, Appendix S1.
pupae are unknown.
Diagnosis: With characters of the erythrocephalus-
group; head orange-red with subtriangular black
9. CREOPHILUS ALBERTISI (FAUVEL)
spot on frons (Fig. 7E), with distinct denticle at hind
(FIGS 1E, 4K, 7E, 28, 31) angles (Fig. 31A, B, vl); antennomeres 7–11 variably
Emus Albertisi Fauvel, 1879: 95, 94 (in key). Type orange-brown to white; right mandible with two dis-
locality: ‘Nouvelle-Guinée, Fly River’; Macleay, 1886: tinct teeth (Fig. 31C); elytra metallic blue-green or
142. violet; abdominal segments VIII–X orange; tergal cha-
Creophilus Albertisi; Bernhauer & Schubert, 1914: etotaxic formula = 4-6-6-2-4-4.
398; Scheerpeltz, 1971: 202.
Creophilus albertisi; Gressitt & Hornabrook, 1977: Description: Measurements (N = 10씹, 10씸). Forebody
26, pl. 5h; Herman, 2001b: 3315. length: 씹 6.3–9.7 mm, 씸 6.8–8.4 mm. See supporting
Table S5 for comparison of ranges of male and female
Type material: Emus Albertisi. Lectotype (here ratios. Head (Fig. 27A, B) orange-red, narrowly black
designated). 씹, ‘Nuova Guinea| Fly River| around mouthparts and antennal fossae, frons with
L.M.D’Albertis 1876–77/ Co Typus/ Museo Civico| di subtriangular black spot usually extending to fronto-
Genova/ FMNH-INS 0000 016 766/ [red] LECTO- clypeal margin, and concealing dorsal tentorial pits
TYPE| Emus| albertisi Fauvel, 1879| designated on vertex or not; very slightly trapezoidal to subquad-
by| D.J. Clarke 2008’ (in MCSN). Paralectotypes (15). rate in large males, suborbicular in females and
All with same collecting data as lectotype and with smaller males; HW/HL = 1.14–1.55; hind angles
label ‘[yellow] PARALECTOTYPE| Emus| albertisi sharply delimited (Fig. 31A); surface dull, with dis-
Fauvel, 1879| designated by| D.J. Clarke 2008’. Five tinctive micropunctate microsculpture and very
specimens with label ‘Museo Civico| di Genova’: shallow indistinct and moderately dense impressions
2씹, ‘Co Typus/ FMNH-INS 0000 016 767’; ‘Co (as in Fig. 9A, B); ventrolateral carina present in both
Typus/ FMNH-INS 0000 016 768’; 3씸, ‘Co Typus/ sexes (Fig. 31B, vl), also visible dorsally (Fig. 31A),
FMNH-INS 0000 016 769’; ‘Co Typus/ FMNH-INS developed into distinct anteriorly projecting denticle;
0000 016 770’; ‘Typus/ Albertisi| Fauvel./ C. Alber- postgenal ridge obsolete medially (Fig. 31B, pgr); eyes
tisi| FvL./ FMNH-INS 0000 016 765’ (in MCSN); five large, protruding (EYL/HL = 0.47–0.68), lateral to
specimens with label ‘Ex-Typis’: 3씹, ‘albertisi| FvL./ dorsolateral in large males; lateral margins of head
R.I.Sc.N.B. 17.479| Coll. et det. A. Fauvel/ FMNH- usually obscured by eye in dorsal view; HL1/HL2
INS 0000 016 760’; ‘Coll. et det. A. Fauvel| Creophi- greater in females than males (씹 = 1.46–2.88,
lus| albertisi| Fauv.| R.I.Sc.N.B. 17.479/ FMNH-INS 씸 = 2.29–3.40); antennae as in Figure 31G, antenno-
0000 016 761’; ‘Coll. et det. A. Fauvel| Creophilus| meres 1–6 black, 7–11 yellowish-white to mottled
Albertisi| Fauv./ R.I.Sc.N.B. 17.479/ FMNH-INS orange-brown (Fig. 1E), 11 elongate, as long or longer
0000 016 763’; 2씸, ‘Coll. et det. A. Fauvel| Creophi- than 9–10 together; mandibles as in Figure 31C, mod-
lus| albertisi| Fauv.| R.I.Sc.N.B. 17.479/ FMNH-INS erately longer than head in large males, shorter than
0000 016 762’; ‘Coll. et det. A. Fauvel| Creophilus| head in females (ML/HL 씹 = 0.78–1.27, 씸 = 0.74–
Albertisi| Fauv./ R.I.Sc.N.B. 17.479/ FMNH-INS 0.92), right mandible with two teeth, T1 absent, T3
0000 016 764’ (in IRSNB); 씸, ‘Emus| albertisi| largest. Thorax and abdomen. Pronotum (Fig. 31F)
Fauvel/ coll. L.W.| Schaufauß/[red] SYNTYPUS| slightly transverse (PW/PL = 1.03–1.26); PL 1.09–
Emus| albertisi Fauvel, 1879| labeled by MNHUB 1.29 ¥ ESL; with basolateral margins slightly emar-
2004/ FMNH-INS 0000 016 759’ (in ZMHB); 3 speci- ginate; hind angles distinct; with sparse peripheral
mens on card: 2씹, FMNH-INS 0000 019 622; FMNH- setae, longer sparse vestiture on lower anterolateral
INS 0000 019 623; 1씸, ‘[Creophilus| albertisi| declivities, and long medially directed basal setae;
Fauvel/ FMNH-INS 0000 019 622’ (in MNHN); 1씹, small transverse basolateral impressions present,
788 D. J. CLARKE
Figure 31. Creophilus albertisi, details of morphology (setae of head, mandibles, antenna, and pronotum omitted). A,
head (large male), dorsal; B, same, ventral; C, mandibles, dorsal (bases omitted); D, apex of median lobe with fully everted
internal sac, right lateral; E, aedeagus with details of inverted internal sac, right lateral (parameral setae omitted) –
arrow indicates dorsoapical projection of median lobe; F, pronotum (large male); G, right antenna, dorsal (only apical setae
shown); H, ventral sclerite of internal sac, ventral; I, female internal genitalia, ventral – arrow indicates thin medial
sclerite of vaginal fold; J, paramere, dorsal. Abbreviations: T2, T3, mandibular teeth; lr, lateral spiculose region;
pgr, postgenal ridge; vl, ventrolateral carina. Additional abbreviations as in Figure 3 (male) and Figure 5 (female).
Scale bars: H = 0.1 mm; rest = 1 mm.
smooth and asetose, with transverse line of tiny middle; wings fully developed, clear yellowish-brown,
punctules; elytra distinctly widened posteriorly, without black spot in medial field between MP3
brightly coloured metallic blue-green or violet and MP4 veins; abdomen with segments III–VII
(Fig. 1E), sparsely and finely setose, without sculp- shining black, VIII–X orange (Fig. 1E); vestiture fine,
ture; humeri sparsely punctate, narrowly glabrous moderately dense; microsculpture extremely fine, pro-
laterally, not callused; disc shallowly impressed in ducing weak iridescent reflection, particularly on
ventral side; tergite VII with well-developed palisade Remarks: Last (1987) recorded Creophilus from New
fringe. Male genitalia and secondary sexual charac- Guinea, but did not list the species, nor did he provide
ters. Sternite IV with densely setose posteromedial any records.
emargination of transverse fold (most obvious in
larger specimens), and sternites V–VII with sparsely
distributed micropores near middle of sternites. 10. CREOPHILUS OCULATUS (FABRICIUS)
Aedeagus as in Figure 31E; median lobe distinctly
yellowish, not produced apically, with minute projec- (FIGS 1C, 3J, 4G, 5D, 7C, 9J, K, 32,
tion dorsoapically (Fig. 31E, arrow), without lateral 33A, D, G, J, M, N, Q, 34)
recurved teeth, with paired apicolateral Staphylinus oculatus Fabricius, 1775: 265. Type local-
sclerites (as) separated from sclerotized median lobe ity: ‘nova Hollandia et Zelandia’ (here restricted to
by membranous strip. Paramere nearly as wide as New Zealand, see below); Goeze, 1777: 724; Fabricius,
median lobe, with very long apicolateral setae 1781: 335 (not seen); Fabricius, 1787: 220; Gmelin,
(Fig. 31J). Internal sac inverted as in Figure 31E, 1790: 2026 (not seen); Fabricius, 1793: 521; Olivier,
everted as in Figure 31D; with ventromedian spicu- 1795:(42): 11, pl. 2, fig. 19; Fabricius, 1801: 592;
lose strip (vr) reduced, shorter than ventral sclerite Turton, 1802: 510; Gravenhorst, 1806: 126; Olivier,
(vs), flanked by very large paired basoventral spicu- 1808: pl.2, fig. 19 (not seen); Boisduval, 1835: 54, pl.
lose regions (Fig. 31D, sr), with additional lateral 9, fig. 1; Erichson, 1839: 352; Dieffenbach, 1843: 273;
spiculose regions (Fig. 31D, lr); ventral sclerite White, 1846: 6; Erichson, 1859 (Fogg translation, not
expanded distally (Fig. 31H); copulatory piece (cp) seen): 312; Murray, 1870: 57, 89; Wakefield, 1873:
completely surrounded by distinctly spiculose mem- 299; Hutton, 1874: 160; Broun, 1876: 265, 271; von
branous sheath (sh). Female internal genitalia. Inter- Kiesenwetter, 1877: 161; von Kiesenwetter & Kirsch,
nal female genitalia as in Figure 31I; vaginal plate 1877: 154 (not seen); Broun, 1880: 107; Broun, 1882:
(vp) small, paired lateral sclerites fused posteriorly to 9; Hudson, 1892: 25, pl. 1, fig.5; Hutton, 1898: 155;
form narrowly sclerotized ring; vaginal fold with Hutton, 1904: 183; Hudson, 1923: 362; 1934: 45, 180;
medial sclerotized strip proximally (Fig. 31I, vf, Zimsen, 1964: 230; Radford, 1981: 185.
arrow), and with diffuse patch of minute teeth dis- Creophilus oculatus; Nordmann, 1837: 23; Mots-
tally. Chaetotaxy. Basiantennal, parasutural 1, and chulsky, 1858a: 49; Gemminger & Harold, 1868: 575;
posterior epipleural macrosetae absent; elytral discal Fauvel, 1875: 56; Reitter, 1880: 166; Fauvel, 1885:
series with three macrosetae; metaventrital mac- 312; Olliff, 1887: 493; Alfken, 1903: 604 (not seen);
roseta absent or undetected; tergal chaetotaxic Walker, 1904: 72, 121; Broun, 1909: 145; Broun, 1910:
formula = 4-6-6-2-4-4, inner lateral macrosetae absent 292; Bernhauer & Schubert, 1914: 399; Brookes,
on tergites VI–VIII, medial macrosetae absent on 1925: 286; Lea, 1925: 229; Steel, 1949: 59, figs 3 and
tergites III and VI. 6; Gourlay, 1950: 185, 202, figs 1 and 2 (pls. 21, 22);
Brookes, 1951: 13, 29; Manson, 1960: 59; Eyles, 1961:
132; Coiffait, 1976: 218; Watt, 1980; Radford, 1981:
Distribution (Fig. 28, circles): Papua New Guinea, 186; Andrews, 1986: 40; Andrews & Gibbs, 1989: 107,
Indonesia: Irian Jaya. Scheerpeltz (1971) notes 113; Kuschel, 1990: 17, 26, 45; Klimaszewski & Watt,
it is common throughout New Guinea and in 1997: 116, figs 4 and 5, p.124, 57; Emberson, 1998: 35;
(unspecified) neighbouring islands, but I have not Ward et al., 1999: 104; Herman, 2001a: 52 (senior
seen any specimens from outside mainland New primary homonym of Staphylinus oculatus O. Müller,
Guinea. 1776: 99); Herman, 2001b: 3325.
Emus oculatus; Fauvel, 1877: 250; Fauvel, 1878a:
248; Fauvel, 1885: 311.
Biology and ecology: Rare in collections; most speci- Creophilus erythrocephalus var. oculatus; Lea,
mens without habitat or collection data. Fifteen col- 1925: 229 (synonym of C. erythrocephalus).
lections are from light traps (mercury vapour, black
light), two from carrion. Habitat: unknown, but prob- Type material: Staphylinus oculatus Fabricius. Lec-
ably intact forest. Altitude: 60–1700 m. Phenology: totype (here designated). 씹, ‘Staphyl. oculatus|
throughout the year. Gressitt & Hornabrook (1977: Fab. Entom.p.265.4/ [red] LECTOTYPE| Staphyli-
26) gave the only known published biological obser- nus| oculatus Fabricius, 1775| designated by| D.J.
vations, noting that C. albertisi ‘occasionally comes to Clarke 2008’ (in BMNH). Specimen with antenno-
light’ and ‘can discharge a peculiar pungent scent meres 4–11 and right metatarsomeres 3–5 missing,
when frightened’. Other biology and life-history char- a smaller pin hole through base of pronotum, and
acteristics are unknown. Larvae and pupae are prothorax glued to mesothorax. Wings are unfolded,
unknown. intact. Paralectotypes (2). 1씹 [incorrectly as 씸 by
790 D. J. CLARKE
Radford 1981], no data but with labels ‘[yellow] and short, sparse vestiture on anterolateral declivi-
PARALECTOTYPE| Staphylinus| oculatus Fabri- ties; elytra uniformly black, disc moderately densely
cius, 1775| designated by| D.J. Clarke 2008’ (in setose, humeral regions glabrous and shining, rug-
BMNH). Specimen missing left antenna, right osely sculptured; wings fully developed (Fig. 33N),
antennomeres 3–11, and right metatarsus; glued to functional, clear black with distinct black spot in
pin on ventral side. Wings are unfolded, intact. 1씸, medial field between MP3 and MP4 veins; mesoven-
no data but with labels ‘[yellow] PARALECTO- tral intercoxal process broadly truncate to convex
TYPE| Staphylinus| oculatus Fabricius, 1775| des- medially (Fig. 33G); metaventrite without posterome-
ignated by| D.J. Clarke 2008/ Creophilus oculatus dial tumescence; protibiae always with outer row of
(Fabricius, 1775) det. D.J. Clarke 2010’ (in ZMUC). long spines (Fig. 33M); last protarsomere much longer
Specimen pinned in a unit tray with label pinned than preceding two together (Fig. 33J); abdomen
in tray: ‘ocula| tus.’ [Fabricius’s handwriting]. All shining and uniformly black; tergite VII with well-
three specimens are pinned through the right developed palisade fringe (Fig. 1C). Male genitalia.
elytron with thick pins typical of the period Aedeagus as in Figure 32C and D; median lobe with
(Andrews & Gibbs, 1989). apex extended to short globular point (Fig. 32I); with
basal and apical subtriangular membranous regions
Other material examined: 1443 specimens. See sup- ventrally (Figs 3J, 32C, I, stm); apicolateral sclerites
porting information, Appendix S1. fused to lateral edge of median orifice (Figs 3J, 32,
as). Paramere shaft distinctly narrower near base
Diagnosis: With characters of the erythrocephalus- than apically (Fig. 32H). Internal sac inverted as in
group; temporal regions and postgena orange-red, Figure 32C and D, partially everted as in Figure 3J.
rest of integument and elytra uniformly pitchy-black Female internal genitalia. Internal female genitalia
(Fig. 1C); apex of antennomere 11 produced into as in Figure 32G; vaginal plate (vp) without median
distinct ridge (Fig. 32B); macropterous (Fig. 33N); sclerotized strip, with paired lateral sclerites (pls),
last protarsomere much longer than preceding two posterolateral areas membranous; vaginal fold (vf)
together (Fig. 33J); elytral discal series with three or strongly and finely rugosely sclerotized (Fig. 32G, vf).
four macrosetae; tergal chaetotaxic formula = 4-6-6-6- Chaetotaxy. Elytral discal series with 3–4 macrosetae;
4-6. metaventrital macroseta present; protrochanter with
two macrosetae, mesotrochanteral macroseta present;
Description: Measurements (N = 10씹, 10씸). Forebody tergal chaetotaxic formula = 4-6-6-6-4-6, medial mac-
length: 씹 5.7–9.9 mm, 씸 5.2–8.0 mm. See supporting rosetae absent from tergite III, inner lateral macrose-
Table S5 for comparison of ranges of male and female tae absent from tergite VII.
ratios. Head. Head (Figs 7C, 33A) black between eyes,
posteriorly including neck, and narrowly around Comparison: Creophilus oculatus may be distin-
mouthparts ventrally, temporal region and postgena guished from all other Creophilus species (except
orange-red, head thus appearing dorsally as ‘black rarely C. huttoni) by the distinct red ‘spots’ behind the
with red spots behind eyes’; moderately to strongly eyes and red postgena. It may be distinguished
trapezoidal, much wider posteriorly; HW/HL = 1.38– from C. huttoni, which has an identical colour
1.67; shining, without distinct microsculpture; eyes pattern (though usually of different colours), by the
small to moderately large (EYL/HL = 0.39–0.57), dor- pitchy black, generally shining cuticle, its fully devel-
solateral, lateral margins of head visible in dorsal oped and functional wings (Fig. 33N, Q), and the
view (not obscured by eye); HL1/HL2 greater in broader obtuse or blunt mesoventral intercoxal
females than males (씹=1.21–2.00, 씸 = 2.00–2.50); process.
antennae as in Figure 32B, antennomeres 1–6 black,
7–11 greyish-black, 11 slightly shorter than 9–10 Distribution (Fig. 34): Many entomologists have con-
together, with apex asymmetrically produced into dis- sidered C. oculatus to be introduced from Australia.
tinct ridge (Fig. 32B, arrow); mandibles as in Herman (2001b) recorded it from the Chatham
Figure 32A, longer than head in large males, shorter Islands, New Zealand, Australia, New Guinea, and
than or subequal to head in females (ML/HL Society Islands (French Polynesia). Coiffait (1976)
씹 = 0.90–1.37, 씸 = 0.86–1.00), right mandible with recorded it from Tahiti (Arue) and only older litera-
three teeth, T3 largest, T1 reduced in smaller speci- ture records it from New Guinea (Fabricius, 1781,
mens. Thorax and abdomen. Pronotum (Figs 32F, 1801; Olivier, 1795); I have seen no specimens from
33D) slightly transverse (PW/PL = 1.04–1.24); moder- either places. Gemminger & Harold (1868) recorded it
ately longer than elytral suture (PL 1.23–1.50 ¥ ESL); from Brazil (an error according to Brookes, 1951).
with basolateral margins very slightly emarginate, Brookes (1951) considered it adventive on the
hind angles indistinct; with sparse peripheral setae Auckland Islands, and it is possible that it did not
Figure 32. Creophilus oculatus, details of morphology (setae of mandibles, antenna, and pronotum omitted). A, man-
dibles, dorsal (bases omitted); B, right antenna, dorsal (only apical setae shown); C, aedeagus with details of inverted
internal sac, ventral; D, same, right lateral (parameral setae omitted); E, ventral sclerite of internal sac, ventral (lower)
and apical (upper); F, pronotum (female); G, female internal genitalia, ventral; H, paramere, dorsal; I, median lobe apex,
ventral. Abbreviations: T1, T2, T3, mandibular teeth. Additional abbreviations as in Figure 3 (male) and Figure 5
(female). Scale bars: E = 0.1 mm; rest = 1 mm.
naturally occur on the New Zealand subantarctic, dispersal throughout the Tasman and South
Chatham, or Kermadec Islands previous to the Pacific Region (K. Wise, unpub. map in Lowe, 1973).
whaling and sealing period of the 19th century, when An abundance of seabird or mammal carrion and
the species could easily have been transported to and other detritus on these islands could have facilitated
between these islands via whaling ships. This is establishment.
consistent with Gourlay’s (1950: 186) account The type locality of C. oculatus was given by Fab-
and previous accounts of ship-mediated insect ricius as ‘nova Hollandia et Zelandia’, the same
792 D. J. CLARKE
Figure 33. New Zealand Creophilus species, details of morphology (setae of heads, pronota, mesoventrites, and legs
omitted). C. oculatus (A, D, G, J, M, N); C. huttoni (B, E, H, O); C. rekohuensis, holotype (C, F, I, K, L, P). A–C, head,
dorsal; D–F, pronotum; G–I, mesoventrite; J, K, protarsus, dorsal; L, M, right protibia, anterior; N–P, left wing, dorsal;
Q, scatter plot of wing length vs. pronotum length as proxy for body size (circles = C. oculatus, squares = C. huttoni,
triangles = C. rekohuensis; black symbols = males, white symbols = females). Abbreviations: CuA, MP3, MP4, wing veins
mentioned in text. Scale bars = 1 mm.
as just one other of his beetle species (a dermestid). Gibbs (1989: 107) noted the likelihood that C. ocula-
It has been historically recorded from (eastern) tus was actually collected aboard the Endeavour: ‘Any
Australia by Fauvel (1877), but Olliff (1887) and Lea such dry or decayed animal materials [brought on
(1925) both doubted its presence there. In reference to board] are likely to have transported carrion beetles
one of few records with specific locality data (Queen- to the chart room table . . .’. It is therefore probably no
sland, Rockhampton), Steel (1949: 57) stated: ‘this is coincidence that the carrion-associated dermestid
the only Australian specimen seen up to the present’. (Dermestes carnivorus F.) has the same type locality.
More recently, Andrews & Gibbs (1989: 107) noted Kuschel (1970: 191), Radford (1981: 158), and
that C. oculatus ‘may have been found in both New Andrews & Gibbs (1989: 110) detail many problems
Zealand and Australia’ during Captain Cook’s H.M.S. with the material collected on the 1769 Endeavour
Endeavour voyage and Kuschel (1990: 17) stated that voyage (and later described by Fabricius), and most
it is ‘an Australian species, but was already in New importantly they indicate where several New Zealand
Zealand at the time of Captain Cook’s visit in 1769’. species had been erroneously described from Austra-
Some ecological works also considered it adventive in lia or South America, and vice versa. It now seems
New Zealand (e.g. Ward et al., 1999). Andrews & most likely that C. oculatus types were either col-
Figure 34. Distribution of all georeferenced specimens of C. oculatus (circles) on the New Zealand mainland (left) and
Chatham Islands archipelago (right). This species is also known from several other offshore islands, including the
subantarctic Auckland and Snares Islands in the south, and the Kermadec Islands in the north east. Rectangle at
southern end of New Zealand indicates area shown in Figure 35.
lected on-board the H.M.S Endeavour or were victims glands (pers. observ.). Kuschel (1990) found it in open
of mixing of material, and that this species is indeed areas (pasture), and on beaches. It is also typically
endemic to New Zealand or the New Zealand subre- found in synanthropic situations and is commonly
gion. More recent literature lists C. oculatus as a New assumed to be a species primarily of open habitats (e.g.
Zealand endemic (Klimaszewski et al., 1996; Leschen Watt, 1980; Kuschel, 1990), but evidently can be found
et al., 2003). in a wide range of habitats. Most specimens have been
taken from carrion or carrion-baited pitfall traps, and
Biology and ecology: Creophilus oculatus is the largest many records are from forest habitat (c. 350 speci-
and one of the most common rove beetles in the New mens). Other specimens were taken at light, in
Zealand Region. It is widely recorded as a typical Malaise traps, window traps, under seaweed, and by
carrion associate, and Gourlay (1950) gives an account many other collecting methods. Habitat: forest of
of maggot predation. When exposed to bright light, all kinds (e.g. Agathis, Nothofagus, mixed podocarp/
individuals more or less immediately attempt to fly, broadleaf), swamp forest, scree and tussock land,
and when threatened discharge an unpleasant- penguin rookeries, pasture, coastal dunes, and
smelling white substance from the abdominal defence beaches. Altitude: sea level to 2440 m.
794 D. J. CLARKE
Phenology: throughout the year, although few records into distinct ridge; brachypterous (Fig. 33O); last pro-
from June–August. Larvae are known and will be tarsomere much longer than preceding two together;
described elsewhere. Pupae are unknown. elytral discal series with two macrosetae; tergal cha-
etotaxic formula = 4-6-6-6-4(6)-6.
Remarks: Zimsen (1964) listed three syntypes for
Staphylinus oculatus Fabricius, two in BMNH and Description: Measurements (N = 10씹, 10씸). Forebody
one then in Kiel (now in ZMUC). Radford (1981: 186) length: 씹 5.5–8.5 mm, 씸 6.0–7.8 mm. See supporting
listed a second label on the lectotype specimen Table S5 for comparison of ranges of male and female
(‘BMNH Staphylinus oculatus’) but this was not ratios. Head. Head (Fig. 33B) between eyes usually
present in 2005 when I examined it. There is now variegated reddish-brown to uniformly dark brown or
ample evidence to correct the type locality to New black, often with diffuse medial lighter patch, or var-
Zealand, following Recommendation 76A.2 (ICZN, iegated patterning (e.g. Fig. 1B), yellowish or almost
1999). Under Article 76.1.1 (ICZN, 1999) the type red (in darker specimens) behind eyes and on
locality of C. oculatus is therefore corrected to ‘New postgena; shorter and narrower in females and sub-
Zealand’. Herman (2001b: 52) determined Staphyli- equal in males to pronotum (HL/PL = 0.6–0.79,
nus oculatus O. Müller, 1776: 99 (Type locality: HW/PW = 0.83–1.12), transverse (HW/HL = 1.29–
Dania) to be a junior primary homonym of C. oculatus 1.63); eyes moderately large (EYL/HL = 0.41–0.55),
Fabricius (ex Staphylinus), but regarded the Müller dorsolateral, lateral margins of head visible in dorsal
name as a nomen dubium and suggested a replace- view (not obscured by eye); HL1/HL2 greater in
ment name was not really needed. females than males (씹 = 1.25–1.83, 씸 = 1.71–2.20);
antennomeres 1–6 reddish-brown, 7–11 lighter
yellowish-brown; mandibles moderately longer than
11. CREOPHILUS HUTTONI (BROUN) STATUS RESUR.
head in large males, subequal to or shorter than head
(FIGS 1B, 5C, 33B, E, H, O, Q, 35) in females (ML/HL 씹 = 0.94–1.22, 씸 = 0.9–1.06),
Staphylinus huttoni Broun, 1880: 108. Type locality: right mandible with three teeth, T3 largest, T1
‘near Dunedin’; Fauvel, 1885: 312; Hutton, 1904: 183; reduced in smaller specimens; left mandible with
Hudson, 1923: 362; Tillyard, 1926: 209, pl. 17, fig. 27, distal tooth projecting from base of mandibular blade.
text-fig. R37; Hudson, 1934: 180; Gourlay, 1950: 186 Thorax and abdomen. Pronotum (Fig. 33E) slightly
(synonym of C. oculatus, attributed to W.O. Steel, transverse (PW/PL = 1.00–1.18), variegated reddish-
p. 202). brown to uniformly dark brown; distinctly longer than
Creophilus huttoni; Walker, 1904: 125; Herman, elytral suture (PL 1.45–1.76 ¥ ESL; elytra reddish
2001b: 3325 (synonym of C. oculatus). brown, moderately densely covered in golden-brown
Creophilus Huttoni; Bernhauer & Schubert, 1914: vestiture; brachypterous, wings short (Fig. 33O, Q),
398. clear yellowish-brown without distinct spot in medial
field, non-functional; mesoventral intermesocoxal
Type material: Staphylinus huttoni Broun. Holotype process moderately to distinctly obtusely angulate
(by monotypy). 씹, ‘200./ [circular with red border] (Fig. 33H), often produced into short subacuminate
Type/ Otago.| – Hutton/ New Zealand.| Broun apex; metaventrite broadly flattened, occasionally
Coll.| Brit. Mus.| 1922–482./ Creophilus| huttoni./ with small posteromedial tumescence near hind
[circular with red border] Holo-| type/ FMNH-INS margin; protibiae always with outer row of spines (as
0000 016 778/ [red] HOLOTYPE| Staphylinus| in Fig. 33M); last protarsomere much longer than
huttoni Broun, 1880| teste D.J. Clarke 2008’ (in preceding two together (as in Fig. 33J); abdomen uni-
BMNH). ‘near Dunedin’ (in Otago) is not on any label formly reddish-brown to brownish-black; tergite VII
of the holotype, but the pronota of other BMNH males with at most poorly developed palisade fringe. Male
do not have ‘indistinct impressions before the middle’ genitalia. Aedeagus indistinguishable from that of C.
as Broun stated, and which are present on the holo- oculatus, see Figure 32C and D. Female genitalia.
type; other specimens in Broun’s collection are Internal female genitalia as in Figure 5C; vaginal
females. plate (vp) without median sclerotized strip, with
paired lateral sclerites (pls), posterolateral aspects
Other material examined: 220 specimens. See sup- membranous; vaginal fold (vf) with single large scler-
porting information, Appendix S1. ite, finely rugosely sculptured. Chaetotaxy. Elytral
discal series with two macrosetae; protrochanter with
Diagnosis: With characters of the erythrocephalus- two macrosetae, mesotrochanter with one macroseta;
group; temporal regions and postgena yellowish to tergal chaetotaxic formula = 4-6-6-6-4(6)-6; medial
reddish-brown, rest of body variegated reddish-brown macrosetae absent from tergite III, inner lateral mac-
to brownish-black; apex of antennomere 11 produced rosetae present or absent on tergite VII.
Figure 35. Distribution of all georeferenced specimens of C. huttoni in southern New Zealand (circles). For location of
map within New Zealand see rectangle in Figure 34.
Variation: Creophilus huttoni varies in colour from Biology and ecology: Creophilus huttoni is an obli-
very light mottled brown with darker patches on the gate beach- and dune-inhabiting species, found
head and pronotum through to uniformly dark beneath drier seaweed, particularly older buried
reddish-brown. The latter specimens occasionally also algal holdfasts high above the strand line. Speci-
lack the abrupt change to yellowish or reddish at the mens have been taken from beach drift, in wrack,
temples (the normal pattern in both C. huttoni and C. supralittoral logs, and under bull kelp, rocks, and
oculatus), but this is rare. The pronotum of this species debris. It is found mainly on sandy beaches, but
is only subtly different from that of C. oculatus, usually seems abundant only where there is a wide supralit-
with straighter sides, and varies somewhat with size. toral zone (pers. observ.). I collected and observed
many specimens live during December 2005 and
Comparison: Creophilus huttoni may co-occur with C. found them to be flightless (see description of obser-
oculatus on sandy beaches where it is found, and no vations of attempted flight in C. rekohuensis, below).
apparent differences between aedeagi of C. huttoni Like the related intertidal species Thinopinus pictus
and C. oculatus specimens were observed. They are (Richards, 1982), the major food source appears to
externally distinguishable by general habitus and be amphipods, and these were abundant above the
(usually) coloration. Creophilus huttoni also has the strand line. Maggots or other larvae were most
following morphometric characters: more quadrate abundant in fresher seaweed lower on the shore
pronotal shape, shortened and narrower elytra, where Cafius was very abundant, but C. huttoni
obtusely angulate mesoventral process (Fig. 33H), rare. I intercepted one specimen as it was feeding
and reduced, non-functional wings (Fig. 33O, Q). on an amphipod, and fed several specimens amphi-
Figure 33Q clearly illustrates a discrete difference in pods for more than a week. Larvae were found in
wing development between these two species. hollowed-out sand galleries, particularly beneath
algal holdfasts above the strandline (pers. observ.).
Distribution (Fig. 35): New Zealand (South Island): A few specimens discharged an unpleasant smelling
Otago Peninsula, south to Stewart Island and sur- white substance from the abdominal defence
rounding islands. One specimen in CMNZ (FMNH- glands when threatened. Phenology: January–April,
INS 0000 017 658) is labelled from Kaitaia (North August–December. Larvae are known and will be
Island: ND), but is probably mislabelled. described elsewhere. Pupae are unknown.
796 D. J. CLARKE
Remarks: The pronotal ‘impressions’ noted by Broun Res [44 °19′32″S 176 °15′46″W]| 22.xi.1992| J.W.
(1880) are not unique to C. huttoni, so are diagnosti- Early/ Under rock on| Sarcocornia-| Disphyma papil-
cally useless. Gourlay (1950: 202) acknowledged W. O. latum| sward/ AMNZ 11132| AUCKLAND|
Steel for the synonymy of C. huttoni with C. oculatus, MUSEUM| NEW ZEALAND’ (in AMNZ). 1씸, on point,
and Steel’s advice is likely to have been based on a ‘Rabbit Is [44 °14′28″S 176 °16′55″W]| Chatham Is.|
comparison of aedeagi, which are qualitatively indis- M.A. Ritchie| 7.VIII.68/ FMNH-INS 0000 017 672’ (in
tinguishable. In addition to characters cited above, the CMNZ). 1씸, missing front and hind legs, terminal
restricted distribution of C. huttoni and the unusual abdominal segments and left elytra on card, ‘on
beach-dwelling ecology supports the reinstatement of ground| South East I. [Rangatira I.]| Chatham Is.| 27
C. huttoni to a valid species. Future work on this Dec. 1937| E.G. Turbott/ AMNZ 11133| AUCKLAND|
species should incorporate molecular data and the MUSEUM| NEW ZEALAND’ (in AMNZ). 1씹, ‘NEW
powerful tools of geometric morphometrics, and should ZEALAND, CI| Rangatira| 27–30.xi.1992| J.W. Early
attempt captive rearing and breeding experiments. R.M. Emberson| P. Syrett pitfall traps/ FMNH-INS
0000 012 564’ (in LUNZ). 1씸, ‘NEW ZEALAND, CI|
Rangatira| 28.xi.1992| P. Syrett| under log/ FMNH-
12. CREOPHILUS REKOHUENSIS CLARKE SP. NOV.
INS 0000 017 639’ (in LUNZ). 1씹, not dissected, in
(FIGS 1D, 33C, F, I, K, L, P, Q, 36, 37) 95% alcohol, ‘NEW ZEALAND: Chatham Is.| Ran-
Type material: Holotype. 씹, ‘NEW ZEALAND: gatira I. Nature Res., Top Bush,| Old Forest Rd. Trk.,
Chatham| Is. Rangatira I. Nature| Res., Woolshed 90 m, 44 °20.87′S,| 176 °10.5′W, 9–14.ii.2006, coastal|
Bush,| Woolshed Bush Trk.,30 m,| 44 °20.45′S, broadleaf forest (Myrsine-| Pseudopanax-Coprosma);|
176 °10.58′W,/ 9–14.ii.2006, coastal| broadleaf forest FMHD#2006–048, carrion traps| (squid; live x5), D.J.
(Myrsine-| Pseudopanax-Coprosma);| seabird burrow Clarke & M.| Renner; site DC0017, 95% ethanol|
entrance &/ ground at night,| D.J. Clarke & M. FIELD MUSEUM NAT. HIST./ FMNH-INS
Renner;| site DC0015, 95%ethanol| FIELD MUS. 0000 019 461’ (in FMNH). 1씹, macerated, in 70%
NAT. HIST./ FMNH-INS 0000 019 462/ HOLOTYPE| alcohol, dissected, both wings dissected, ‘NEW
Creophilus| rekohuensis Clarke sp. nov.| designated ZEALAND, CI| South East Island [Rangatira I.],|
by D.J. Clarke 2008’ (in LUNZ). Left wing dissected Kokapu Creek [44 °20′50″S 176 °10′10″W]|
and mounted on card, aedeagus and genital segment 15.i.1997 J.W.M Marris| under rock by creek/ FMNH-
dissected, stored in glycerin in genitalia vial mounted INS 0000 012 566’ (in LUNZ). 1씹, aedeagus not dis-
with specimen. Paratypes (22). Unless otherwise sected, ‘NEW ZEALAND, CI| South East I.,
stated, the left wing, aedeagus and genital segments of [Rangatira I.]| Woolshed Bush [44 °20′S 176 °10′W]|
males are dissected and mounted with the speci- 13–17.i.1997| R.M. Emberson, J.W. Marris/ pitfall
men. All pinned specimens and alcohol lots with traps in coastal| broadleaf forest/ FMNH-INS
label ‘PARATYPE| Creophilus| rekohuensis Clarke 0000 012 562’ (in LUNZ). 1씹, same as preceding speci-
sp. nov.| designated by D.J. Clarke 2008’: 6 specimens, men but with date ‘19–21.i.1998’ [county as ‘CH’] and
same data as holotype: 3씹, 1씸, pinned, [씹’s, ‘FMNH-INS 0000 012 563’ (in LUNZ). 1씹, ‘NEW
FMNH-INS 0000 019 463–465]; [씸, FMNH-INS ZEALAND, CI| South East I., Woolshed| Bush-
0000 019 466]; 2 specimens in 70% alcohol, 1씹, Whalers Bay Track [44 °20′22″S 176 °10′28″W]|
genitalia dissected, with specimen, ‘FMNH-INS 18.i.1997 J.W.M. Marris| on trees and logs at night/
0000 019 467’; 1씸, genitalia dissected, with specimen, FMNH-INS 0000 012 559’ (in LUNZ). 1씹, ‘NEW
‘FMNH-INS 0000 019 468’ (in FMNH; 1씹 in NZAC). ZEALAND, CH| Star Keys [44 °15′S 176 °0′W]|
1씸, on card, ‘Chatham| Islds./ A.E. Brookes| 23.i.1998| R.M. Emberson| under rocks/ FMNH-INS
Collection/ FMNH-INS 0000 012 565’ (in NZAC). 1씸, 0000 012 558’ (in LUNZ).
leg segments on card ‘Mangere I. [44 °15′S 176 °18′W]|
Chatham Islands.| 4–21 Jan., 1924.| Coll. C. Lindsay/
Description of type locality: Woolshed Bush, northern
Staphylinus sp./ FMNH-INS 0000 017 670’ (in CMNZ).
end of island, < 200 m south of Department of Con-
2 specimens, ‘NEW ZEALAND, CH| Mangere I
servation Hut, along track. Habitat: sparse
[44 °15′S 176 °18′W]| 28.x.1993| R.M. Emberson| in
regenerating coastal broadleaf forest with mixed
petrel burrow entrance’: 씹, right mesotarsal claw on
Myrsine, Pseudopanax, and Coprosma trees domi-
card ‘FMNH-INS 0000 012 560’; 씸, ‘FMNH-INS
nant. Understorey sparse to open, ground with high
0000 012 561’ (in LUNZ). 1씸, macerated, in 70%
density of diving petrel burrows, leaf litter scant,
alcohol, both wings dissected, ‘NEW ZEALAND, CH|
ground layer mostly compacted soil.
Pitt Island,| Preece Covenant Bush [44 °16′55″S
176 °10′45″W]| 18.i.1998 J.W.M. Marris| under
rotten log/ FMNH-INS 0000 012 567’ (in LUNZ). 1씸, Diagnosis: With characters of the erythrocephalus-
‘CHATHAM IS, N.Z.| Pitt Island, 100 m| Rangiauria group; postgena pale yellowish to reddish-brown,
Figure 36. Creophilus rekohuensis, details of morphology (A, B, D–H, J, K = holotype; setae of mandibles, antenna,
pronotum, and legs omitted). A, mandibles, dorsal (bases omitted); B, right antenna, dorsal (only apical setae shown); C,
ventral sclerite of internal sac, ventral (lower) and apical (upper); D, paramere, dorsal; E, aedeagus with details of
inverted internal sac, ventral; F, same, right lateral (parameral setae omitted); G, protarsus, dorsal; H, pronotum; I,
female internal genitalia, ventral; J, right protibia, anterior; K, median lobe apex, ventral. Abbreviations: T1, T2, T3,
mandibular teeth. Additional abbreviations as in Figure 3 (male) and Figure 5 (female). Scale bars: C = 0.1 mm;
rest = 1 mm.
distinct from blackish-brown of rest of body; Description: Measurements (N = 10씹, 8씸). Forebody
apex of antennomere 11 asymmetrically produced length: 씹 5.3–6.8 mm, 씸 5.6–6.2 mm. See supporting
(Fig. 36B); brachypterous (Fig. 33P, Q); last protar- Table S5 for comparison of ranges of male and female
somere distinctly shorter than preceding two ratios. Head. Head (Fig. 33C) brownish-black dorsally
together (Fig. 36G); elytral discal series with two (Fig. 1D), postgena yellowish to reddish-brown;
macrosetae; tergal chaetotaxic formula = 4-6-6-6- strongly trapezoidal in both sexes, much wider pos-
4(6)-6. teriorly; HW/HL = 1.44–1.63; hind angles distinctly
798 D. J. CLARKE
rounded; shining, without distinct microsculpture; rosetae, protrochanter with two macrosetae, mesotro-
ventrolateral carina absent in all males seen; eyes chanter with one macroseta; tergal chaetotaxic
moderately large (EYL/HL = 0.41–0.50), dorsolateral, formula = 4-6-6-6-4(6)-6; medial macrosetae absent
lateral margins of head visible in dorsal view (not from tergite III, inner lateral macrosetae present or
obscured by eye); HL1/HL2 slightly larger in females absent on tergite VII.
than males, or subequal (씹 = 1.38–2.20, 씸 = 1.67–
2.17); antennae as in Figure 36B, subclavate, Variation: Several paratypes have the transverse
antennomeres 1–6 brown-black, 7–11 lighter basal ridge of tergites III–V variously effaced or very
reddish-brown, 11 distinctive, without apical ridge, faint. A few specimens are teratological, having
asymmetrically produced to anterior subconical point, deformed heads or pronota.
much shorter than 9–10 together; each apical pair of
setae close together on anterior subconical point; Comparison: Emberson (1998) recorded C. rekohuen-
mandibles as in Figure 36A, subequal to head in sis as a new, undescribed species but noted it may be
males and females (ML/HL 씹 = 1.00–1.06, 씸 = 0.88– identical to C. huttoni. It is best distinguished from
1.12), right mandible with three teeth, T3 largest, T1 other New Zealand species by the strongly trapezoidal
and T2 usually reduced; left mandible with distal head, asymmetrically produced terminal antenno-
tooth strongly projecting from base of mandibular mere (Fig. 36B), and unusually shortened terminal
blade, ventral basolateral ridge weakly developed. protarsomeres (Fig. 36G; unique in Creophilus). Two
Thorax and abdomen. Pronotum (Figs 33F, 36H) further significant differences between C. rekohuensis
slightly transverse (PW/PL = 1.07–1.19); distinctly and all other Creophilus species are (1) apparent
longer than elytral suture (PL 1.59–1.87 ¥ ESL); with sexual monomorphism, and (2) the acutely angulate
basolateral margins very slightly emarginate, hind mesoventral intercoxal process. Although the sample
angles indistinct; disc with sparse peripheral setae size is small, available males and females were rather
and short, sparse vestiture on anterolateral declivi- consistent in size.
ties; elytra subquadrate, width subequal to or slightly
narrower than pronotum, uniformly blackish-brown, Distribution (Fig. 37): Chatham Islands (New
sparsely setose, and rugosely sculptured, especially Zealand): Pitt, Rangatira, Mangere, Star Keys, and
on humeri; brachypterous, wings short (Fig. 33P, Q), Rabbit Islands. Extensive carrion trapping on Pitt
non-functional, clear yellowish-brown, without black and Chatham Islands (27 January to 6 March 2006)
spot in medial field between MP3 and MP4 veins; was unsuccessful in collecting this species. Remaining
mesoventral intercoxal process acutely angulate, seabird nesting colonies are the most likely habitat,
usually distinctly V-shaped (Fig. 33I); metaventrite particularly in large areas of relatively intact native
broadly flattened, with small posteromedial tumes- vegetation such as Tuku Nature Reserve, where there
cence near hind margin; protibiae (Fig. 36J) fre- is also rodent control around taiko (Pterodroma
quently without outer row of spines, when present magentae Giglioli and Salvadori) burrow groups.
with 1–3 very reduced tubercles (much shorter than
accompanying bristles), and rarely with equal Biology and ecology: Collection data indicate C. reko-
number on both protibiae; last protarsomere subequal huensis is nocturnal and active on the ground, but
in length to preceding two together, or shorter rarely found at carrion (only one specimen). However,
(Fig. 36G); abdomen blackish-brown, each segment as far as I know carrion trapping had not been
lighter apically (Fig. 1D); vestiture sparse, brownish- attempted before in the Chatham Islands. Trapping
black; tergite VII with at most poorly developed pali- on Rangatira Island was thwarted by large numbers
sade fringe. Male genitalia. Aedeagus as in of raphidophorid crickets that probably deterred the
Figure 36E and F; with basal and apical subtriangu- beetles, and also quickly consumed much of the bait,
lar membranous regions ventrally (Fig. 36E, K, stm), thereby reducing its effectiveness as an attractant.
with paired apicolateral sclerites (as) fused to lateral Further trapping should utilize much larger and
edge of median orifice; with narrowly acuminate apex better protected baits. Habitat: coastal broadleaf
(Fig. 36K). Paramere as in Figure 36D, usually with forest (mixed Myrsine–Pseudopanax–Coprosma) and
sides subparallel, widest before apex. Internal sac coastal herb fields (Sarcocornia–Disphyma). Altitude:
inverted as in Figure 36E and F. Female internal 30–100 m. Phenology: January–February, August,
genitalia. Internal female genitalia as in Figure 36I; October–December. Light exposure induces a flight
vaginal plate (vp) with paired lateral sclerites (pls), response in which the wings are extended while
with posterolateral aspects membranous; vaginal fold running (pers. observ.). The shortened wings are,
(vf) with single narrow sclerite dorsal to apex of however, non-functional. Wing folding is aided by
vaginal plate (Fig. 36I, vf). Chaetotaxy. Macrosetae repeated curling and telescoping of the abdomen,
unusually long; elytral discal series with two mac- which brings the apex of the seventh tergite into
Chathams-endemic flightless beetles (McGuinness,

2001), its rarity, probable vulnerability to rodent pre-
dation, and possible restriction to the smaller islands
suggests it may require formal conservation status.
Etymology: The species epithet ‘rekohuensis’ is

derived from a combination of Rekohu and the Latin
adjectival suffix -ensis, meaning ‘belonging to’, in ref-
erence to the species being endemic to the Chatham
Islands. Rekohu is the moriori name for Chatham
Island, meaning ‘misty skies’, and refers to the seem-
ingly perennial cloud cover shrouding these islands.
This name was chosen in honour of the moriori
people, the original inhabitants of the Chatham
Islands.
Remarks: The earliest record for C. rekohuensis is a

female specimen collected by C. Lindsay on Mangere
Island in 1924, and it was recorded by Brookes (1925)
as one of two possibly new species of Staphylinus,
along with Quedius antipodum.
ACKNOWLEDGEMENTS
I thank Margaret Thayer and Al Newton for their
many and varied contributions to this project, and
for their patience as I completed it. I am grateful to
all the curators of institutions mentioned in the text
for generously loaning me specimens and many
types for this study, and also for their patience as I
completed it. Roger Booth and Sharon Shute
(BMNH), Alexey Gussakov (ZMUM), Pudji Aswari
(MBBJ), Alexey Solodovnikov (ZMUC), and Mark
Ero (National Agricultural Insect Collection) gener-
ously provided details of some types and other speci-
mens I was unable to study. I also thank Margaret
Thayer for planning and successfully executing
Figure 37. Distribution of georeferenced specimens of C. several field trips, thus facilitating collection of
rekohuensis in the Chatham Islands, New Zealand many Creophilus, and for her tolerance of my per-
(circles). The four largest islands from west to east are sistent desire to collect C. huttoni in New Zealand.
Little Mangere, Mangere, Pitt, and Rangatira. The record For providing logistical support for fieldwork in the
off the north-west coast of Pitt Island is from the tiny Chathams, I sincerely thank Hilary Aikman, Paul
Rabbit Island. This species is also known from Star Keys,
Gasson, and Collin Miskelly (DoC, Chathams Area
14.5 km east of Pitt Island. For location of Chatham
staff). Matthew A. M. Renner and I are very grate-
Islands see Figure 34.
ful to Chatham Islands residents Bill & Dianne
Gregory-Hunt, Johnny Kamo, James Moffet, Abe
contact with the wing surface. Usually several folding Nielsen, Alfred Preece, and Raana Tuuta for access
attempts are made, but even then at least one wing to their land and/or for hospitality, and to the
often remains partly extended. Specimens were Chatham Islands Conservation Board and the New
observed feeding on maggots supplied to them with Zealand Department of Conservation for permission
rotting fish. A few specimens discharged an unpleas- to collect (permit WE/205/res and nature reserve
ant smelling white substance from the abdomen when entry permits 1/06-130 & 131 and 1/06-135 & 136).
threatened. Fieldwork was supported by grants from The Mar-
This new species is morphologically and ecologically shall Field Fund (FMNH) and the J. S. Watson Con-
unique within Creophilus and more detailed servation Trust (Royal Forest and Bird Protection
ecological studies are warranted. Like several other Society, New Zealand). Last but not least, I thank
800 D. J. CLARKE
Matthew A. M. Renner for outstanding assistance Archer MS. 2004. Annual variation in arrival and departure
and companionship on the Chathams, and particu- times of carrion insects at carcasses: implications for suc-
larly for his tolerance of decaying fish heads (!), and cession studies in forensic entomology. Australian Journal
willingness to dig our 4WD out of the swamp. of Zoology 51: 569–576.
Michael Zubich provided assistance with databasing; Arnett RH, Jr, Samuelson GA, Nishida GM. 1993. The
Betty Strack supervised SEM work; and Yvonne insect and spider collections of the world, 2nd edn. Gaines-
Matos, Demonica Jones, and Kathryn Dirks assisted ville, FL: Sandhill Crane Press, Inc.
Asenjo A, Clarke DJ. 2007. Scientific note: first record of
with sample sorting. Margaret Thayer, Alexey
Creophilus maxillosus (L, 1758) (Staphylinidae) for Peru.
Solodovnikov, and Al Newton commented on and
The Coleopterists Bulletin 61: 551.
therefore greatly improved earlier versions of the
Ashe JS. 1984. Generic revision of the subtribe Gyrophaenina
manuscript, and Paul Brinkman is cordially
(Coleoptera: Staphylinidae: Aleocharinae) with a review of
thanked for his constant encouragement as I com- the described subgenera and major features of evolution.
pleted the project (one more month!). This work was Quaestiones Entomologicae 20: 129–349.
supported by US National Science Foundation PEET Ashe JS. 1986. Structural features and phylogenetic relation-
grant DEB-0118749 to Margaret Thayer and Al ships among larvae of genera of gyrophaenine staphylinids
Newton, and the manuscript was completed while (Coleoptera: Staphylinidae: Aleocharinae). Fieldiana:
supported by a Field Museum Lester Armour Zoology (N.S.) 30: 1–60.
Graduate Fellowship. Ashe JS. 2000. Mouthpart structure of Stylogymnusa suban-
tarctica Hammond, 1975 (Coleoptera: Staphylinidae: Aleo-
charinae) with a reanalysis of the phylogenetic position of
the genus. Zoological Journal of the Linnean Society 130:
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APPENDIX 1
MORPHOLOGICAL CHARACTER MATRIX
External Structures (1–48) Male Genitalia (49–77) Female Genitalia (78–91) Chaetotaxy (92–130)
+ ++ + + + + +
10 20 30 40 50 60 70 80 90 100 110 120 130
Staphylinus caesareus 0400000000 0000100-10 0000030102 0000000000 1000000001 100000-000 00--100000 -----00000 0000001101 1011000010 11101*1000 1000000000 1110***110
Creophilus albertisi 1312211131 0011013102 0111101102 0010001011 1001100012 1001110001 1100011211 1010010010 1300221121 1110001100 0110212001 1000000101 1011110000
Creophilus imitator 1012211131 0011013102 0111101102 0010001011 1001100012 1001110111 1101011211 1010011010 1210211121 1110001100 0011212101 1000000111 1101110000
Creophilus erythrocephalus 1013011131 0011013100 0111101102 0010001011 1001100012 1001110111 1101011211 1010011010 1210211121 1111001100 0111212101 1000000101 111111*010
Creophilus lanio 1013011131 0011113100 0111101102 0010001011 1001100012 1001110111 1101011211 1010011010 1110211121 1111001100 0110212001 1000000101 1111111010
Creophilus oculatus 1113011131 0011113100 0111101102 0010001011 1001100012 1010011111 1101011211 1012011010 1110211121 1111001100 0111212101 1100100101 1111111010
Creophilus huttoni 1113011131 0011113100 0111121102 0010101011 1001100012 1010011111 1101011211 1012011010 1110211121 1111001100 0111112101 1100100101 1111111*10
Creophilus rekohuensis 11130?1130 0011113100 ?111121102 0010101011 1001100012 1010011111 1101011211 1012011010 1110211121 1111001100 0111112101 1100100101 1111111*10
Creophilus m. maxillosus 0411001132 0011111001 1111101102 0011011011 1001001012 1100010111 1101011111 1100110010 1011101121 0111001101 0111222001 0001000111 1111110*10
Creophilus m. villosus 0411001132 0011111001 1111101102 0011011011 1001001012 1100010111 1101011111 1101110010 1011101121 0111001101 0111222001 0001000111 1111110*10
Creophilus incanus 0411001131 0011111001 1111101102 0011011011 1001001012 1100010111 1101011111 1100110010 1011101121 0111001101 0011222001 0001000111 1011110010
Creophilus variegatus 0411001131 0011113001 1111101102 0011011010 1001001012 1100111111 0101001111 1000010000 1010101121 1110001101 0101222001 0000000111 111111*110
Creophilus flavipennis 0411001132 0011111001 1111101102 0011011011 1001001012 1100010111 1101011111 1100110010 1011141121 0111001101 0111222001 2001000111 1111111010
Creophilus galapagensis 0411001132 0011111001 1111101102 0010011011 1001001012 1100010111 1101011111 1001010010 1011101121 011*001101 0*002*2001 0001000-00 0000000000
Liusus hilleri 0413101020 0001113000 0101011111 1020000111 1001010012 1010010111 1111001111 1020010010 0010101121 1111111100 01111#1000 2000000111 1111111010
Liusus humeralis 0413101020 0001113000 0101011111 1020000111 1001010012 1010011111 1111001111 1020010010 0010101121 1111111100 01111#1000 2000000111 1111111010
Hadropinus fossor 0413101020 1100102-00 0101011102 0030200201 1011000012 1100010111 1101001111 0000010110 0000100121 1110100100 10*?0??011 3112010100 00111110?0
Thinopinus pictus 0213100020 1100102-00 0*00011100 0100200201 0111000012 1100110?11 1100000101 00?0010011 2000120100 0101001100 1000000011 0112211-00 0000000000
Hadrotes crassus 0413001020 0001102-00 0101011101 0020200111 1001010012 1100010111 1111001111 1020010010 0010101121 1110100100 0110022011 2001000111 11111111?1
Hadrotes wakefieldi 0403201010 0001002-00 0101011002 0020000001 1001100122 1100110002 10-0000000 -----00010 0000231010 0011001100 0110101001 1000000100 0011111110
Xanthopygus xanthopygus 0410300000 0001002-10 0110020002 0000000000 1001100000 0100010000 00-0100000 -----00000 0000001100 0111001110 0110010000 1000000-00 0000000000
Dinothenarus badipes 0400000000 0001002-10 0000010002 0000100000 1000000000 010000-000 00-0100000 -----00000 0000001101 0011001010 1110121000 1000000000 1110000010
Platydracus maculosus 0400000010 0001111010 0000030002 0000000000 1000000000 000100-111 00-0100000 -----00000 0000230010 0011001110 1110121000 1000000000 1110111110
Ontholestes cingulatus 0400000022 0001114010 0000030002 0000000000 1000000000 000100-011 00-0100001 00???00000 0000001100 0011001110 1110111000 1100000000 1110111110
0/1 = *; 1/2 = #; autapomorphies indicated with a + sign at column head.
PHYLOGENETIC UTILITY OF CHARACTERS IN CREOPHILUS
811
812 D. J. CLARKE
SUPPORTING INFORMATION
Additional Supporting Information may be found in the online version of this article:
Appendix S1. Other material examined.
Table S1. Bremer support from each character system for each node in Figures 11–12.
Table S2. Unambiguous synapomorphies from each character system for each node in Figures
11–12.
Table S3. Uniquely optimized synapomorphies from each character system for each node in Figures
11–12.
Table S4. Measurement ratios for species in the C. maxillosus species-group.
Table S5. Measurement ratios for species in the C. erythrocephalus species-group.
Please note: Wiley-Blackwell are not responsible for the content or functionality of any supporting materials
supplied by the authors. Any queries (other than missing material) should be directed to the corresponding
author for the article.

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Zoological Journal of the Linnean Society, 2011, 163, 723–812.

Testing the phylogenetic utility of morphological

INTRODUCTION absence of such data, evolutionary hypotheses and

Figure 6. Scheme of taxonomically and phylogenetically

(Fig. 1F–O); (1) bright orange-red or yellowish,

J). In C. flavipennis (scored as state 1) most MALE GENITALIA (MALE, M)

Table 1. Tree statistics for separate and combined phylogenetic analyses

Data No. of Total no. No. of congruent

S 44 (36) 3 91 91.82 0.71 0.71 0.89 0.89 17 16 (94)

including them here is a much better supported

uting 52% of the total PBS, 42% of the total PSYN,

ences in partition size changed the relative contribu-

group): support contributed by structural and male

while structural characters are clearly superior to

character systems and relative female PUNIQ is even

that some female genital characters are strong

hypotheses of primary homology.

netic signal in the structural and male genital charac-

shown). Nevertheless, two major features of the

support analyses are 1) the high proportion of total

tion of hidden support (HPBS, HPSYN, and HPUNIQ)

contributed by chaetotaxy. Approximately 74% of the

and HPUNIQ (Table 2). Chaetotaxy contributed by far

the greatest amount of hidden support: 62% of HPBS,

of the 18 nodes in the simultaneous analysis have

17) are characterized by zero or negative hidden syna-

contrast between non-chaetotaxy and chaetotaxy

HPSYN): structural characters contributed positive

male genitalia contributed positive HPBS to 11 nodes,

but eight of these (73%) have zero HPSYN and female

genitalia contributed positive HPBS to six nodes, but

female 8/0/1/0 16 C. flavipennis

(B) Creophilus complex 4 40 55 57 61 66 79 94 103 128

8 9 10 13 23 25 26 37 66 81 105 107 16 4 2 1 C. flavipennis

KEY TO ADULTS OF CREOPHILUS SPECIES

CREOPHILUS MAXILLOSUS SPECIES-GROUP Diagnosis: Head black or brownish-black; antenno-

Creophilus maxillosus var. subfasciatus Sharp.

Description: Measurements (N = 10씹, 10씸). Forebody

data). Phenology: February–December. Vinson

Remarks: A female specimen from Réunion in Bern-

3. CREOPHILUS GALAPAGENSIS CLARKE SP. NOV.

humeral macroseta usually absent, if present then

Variation: The extent of abdominal vestiture pattern-

Comparison: Creophilus galapagensis superficially

Figure 22. Distribution of all georeferenced specimens of C. flavipennis (circles).

other species. In describing Emus insularis, Fauvel 5. CREOPHILUS VARIEGATUS MANNERHEIM

Description: Measurements (N = 10씹, 10씸). Forebody Comparison: Creophilus erythrocephalus is most

Description: Measurements (N = 10씹, 10씸). Forebody

Chathams-endemic flightless beetles (McGuinness,

Etymology: The species epithet ‘rekohuensis’ is

Remarks: The earliest record for C. rekohuensis is a

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