Phylogenetic placement of Phaenocephalus Wollaston (Coleoptera: Phaenocephalidae & Phalacridae)

5 Pages·1991·2.2 MB·

by  J Pakaluk

PROC. ENTOMOL. SOC. WASH.
93(2), 1991, pp. 317-321
PHYLOGENETIC PLACEMENT OF PHAENOCEPHALUS WOLLASTON
(COLEOPTERA: PHAENOCEPHALIDAE & PHALACRIDAE)
James Pakaluk
Snow Entomological Museum, University ofKansas, Lawrence, Kansas 66045.
Abstract.—T\it taxonomic history of the family Phaenocephalidae is reviewed. This
family currently consists of two species, Phaenocephalus castaneus Wollaston and P.
coomani Paulian, described from Japan and Viet Nam respectively. The genus is rede-
scribed and selected structures are illustrated. The relationship ofthis enigmatic family
to other beetles has been disputed for years. The uncertainty ofits phylogenetic position
has been due, in part, to the fact that type material has been unavailable for study for
forty years or more. After examining types in collections in London and Paris it seems
that these two species, previously attributed to a separate family, should be placed in the
Phalacridae.
Key Words: Coleoptera, Cucuioidea, Phaenocephalidae, Phalacridae
The Phaenocephalidae is one of the it was most similar to Corylophus Stephens
smallest families of Coleoptera, with only but grouped Phaenocephalus with Sacium
two species described, as well as one ofthe LeConteandMicrostagetusWollastonbased
most enigmatic. Ever since this family was upon their 1 1-segmented antennae. Mat-
described nearly 100 years ago there have thews (1899) established a new family,
been fewpublishedreportsforthesebeetles. Phaenocephalidae, for Wollaston's species
While studying types and other material of and provided a more detailed description
Cucujoidea recently in London and Paris, I and some illustrations. Both authors em-
was able to locate specimens of Phaenoce- phasized the large, deflexed head and an-
phalidae that have likely been unavailable teriorly emarginate edge of the pronotum
for study for forty years or more. Phaeno- in Phaenocephalus. Matthews, however,
cephalus was not listed in the generic list of thoughtthesefeatures,aswellasothers,were
Coleoptera at The Natural History Muse- sufficient to exclude this species from Cor-
um, London, and the material at the Mu- ylophidae. He speculated that Phaenoce-
seum National d'Histoire Naturelle, Paris phalidae was a link between Corylophidae
was among Paulian's dissections of Cory- and Silphidae.
lophidae which were until recently consid- Paulian (1950) described another species
ered lost ordestroyed. It seems appropriate oiPhaenocephalus, P. coomani, from Viet
tonowreviewthetaxonomichistoryofthese Nam and illustrated a few structures. In a
beetles, redescribe the genus, illustrate im- footnote he indicated that he had seen ad-
portant structural features, and discuss the ditional material ofan undetermined spe-
phylogenetic position ofthis family. cieso{Phaenocephalusfrom Sumatra; Iwas
Wollaston (1873) described Phaenoceph- unable to locate this material. I did find,
alus castaneus as a corylophid based upon however, a specimen labelled as Phaeno-
a single specimen from Japan. He thought cephalus sp. from Viet Nam, but this is re-
318 PROCEEDINGS OFTHE ENTOMOLOGICALSOCIETY OFWASHINGTON
ally a species ofOrthoperus Stephens (Cor- available. Thistypeconsistsoftheheadand
ylophidae). prothorax; the rest ofthe specimen appears
There are few subsequent references to tobelost. Thisislikelya syntype since Pau-
thisfamily,andthesewerepresumablymade lianliststhe sizeofthisspeciesas 1.15-1.20
mm
without referringto type material. Crowson long, suggesting more than one speci-
(1955) treated Phaenocephalidae as Cole- men was used for his description. I am not
optera incertaesedis, Lawrence (1982) con- designating a lectotype with the hope that
sidered this family synonymous with Phal- the remaining type material for this species
acridae, and Sasaji (1985) included a will eventually be located.
photograph of Phaenocephalus castaneus
Redescription of
and treated it as a distinct family ofCucu-
Phaenocephalus Wollaston
joidea.
The quality ofpreserved material forthis The following description is based upon
family is extremely poor. Wollaston's type the material listed above. Certain features,
o^Phaenocephaluscastaneusisvirtuallylost; such as total body length, were taken from
the only parts that I could find were the descriptions. Mostdetailsaredescribedfrom
labium and a maxilla mounted in balsam the specimen of the putative P. castaneus
on an acetate card. These structures agree that is in my collection. Every eifort was
with Matthews's (1899) illustrations. All of made to compare these features to previ-
Paulian's material, including the type ofP. ously published descriptions and illustra-
coomani, were poorly mounted on slides tions,aswellasthepreservedmateriallisted
and still in glycerin jelly. These specimens above. It is possible that this description is
have been removed from their slides and inaccurate since the type ofthe type species
transferred to glycerin-filled microvials. ofthis genus is almost entirely lost. In gen-
Therearefewfeaturesthatcanbecompared eral, this agrees with Matthews's (1899) de-
to Wollaston's or Matthews's descriptions scription,exceptthatthemesocoxalcavities
ofP. castaneus. are clearly closed and the ventrites are sub-
I have examined the following material equal in length.
ofPhaenocephalus: Description.—Length 1.15-1.70 mm.
P. castaneus.—The labium and a maxilla Body ovate, about 1.3x longer than wide,
oftheholotype. Thenumberanddate"759/ light brown, glabrous, shiny; dorsum con-
Aug 5 1885," written by Matthews, is at- vex, venter flat.
tached to the pin. Two slide-mounted spec- Head (Fig. 3) transverse, widest at eyes.
imens from Japan were used for Paulian's Eye on antero-lateral margin of cranium.
(1950) monograph. One specimen is rep- Fronto-clypeal suture absent, clypeus
resented by the head only. Paulian (in litt., strongly declivious anteriorly. Antenna 1 1-
1990) never examined Wollaston's type, so segmented with 3-segmented club, apical
these maynotbeconspecific. The specimen club segment elongate, subequal in length
that Paulian listed from Formosa in the to basal two club segments combined; an-
Grouvelle collection was not found. I have tennal groove indestinct. Labrum (Fig. 4)
an additional specimen from Japan identi- transverse, sclerotized, on different plane
fied by H. Sasaji. Most ofthe illustrations from clypeus; tormae (Fig. 4) elongate,
included in this paper were made from this slightly longer than labrum, fused apically
specimen and compared to previously pub- with labial rods, with mesal arms directed
lished descriptions and illustrations. This anteriorly; epipharyngeal rods (Fig. 4) sub-
specimen is in my collection. equal in length to mesal arms of tormae,
P. coomani.—The slide-mounted type fringed with setae mesally. Mandible (Fig.
described by Paulian was the only material 5)bifidapically; molawell-developed;pros-
VOLUME 93, NUMBER 2 319
thecabroad, fringedwithhairs, withasmall subequal in length, segments 1-3 distinctly
basal tuft of setae. Maxilla (Fig. 6) with lobed, 4 elongate, subequal in length to 1-
4-segmented palp, segment 4 longest; galea 3 combined; claws with large, broad tooth
broad, distigalea with dorsal membranous atbase,empodiumabsent. Scutellumsmall,
area basally, with broad brush of elongate triangular. Elytra about 3.2x longer than
setaeapically; lacinianarrow, bifidapically. pronotum, punctationminute,distinctlyse-
Labium(Fig. 8)with 3-segmentedpalp, seg- riate, with 9 intemeurs, intervals smooth
ments 2 and 3 subequal in length; mentum except forextremely fine, seriate punctures;
transverse, anterolateral angles produced; epipleuron large, strongly angled, complete.
prementum mostly membranous, basal Wing (Fig. 2) with venation reduced, cells
scleritesmall,ligulalarge,divergentapically and a subcubital fleck absent, with 2 anal
with distinct lobes, labial rods elongate, veins, jugal lobe present.
fused apically with tormae. Gular sutures Abdomen with 7 spiracles; 5 ventrites
indistinct. Tentorium reduced, represented subequal in length, first with large, narrow
by vestigial anterior arms only. intercoxal process; femoral lines absent; all
Pronotum (Fig. 1)transverse, about0.4x ventrites free; 2-5 with small, anterolateral
longer than wide, anterior edge with deep internal apodemes. Male genitalia with teg-
U-shaped emargination, posterioredge with men (Fig. 9) elongate, with apical subtrian-
large medial lobe, lateral edges convergent gular sclerite deeply emarginate medially,
anteriorly, with smooth margins. Proster- median lobe (Fig. 10) weakly sclerotized.
num reduced anteriorly; intercoxal process I agree with Lawrence's (1982) decision
narrow,elevated,deflexedapically. Procoxa to synonymize Phaenocephalidae with
globose, with long internal extension, its Phalacridae,althoughhedidthisbasedupon
cavity internally closed, externally widely publishedinformationonly(J. F. Lawrence,
open. Mesostemum between coxae sub- pers. comm.). In general habitus, Phaeno-
equal inwidthto mesocoxal cavity, without cephalus is certainly a typical phalacrid. Its
femoral lines, junction ofmeso- and meta- lackofa fronto-clypeal suture, maxilla with
stemum straight-line type, without internal distinctlaciniaandgalea, broadlyopen pro-
knobs. Mesocoxa globose, trochantin con- coxal cavities, laterally closed mesocoxal
cealed, its cavity laterally closed. Metaster- cavities, 4-4-4 tarsi distinctly lobed with
num twice as wide as long, without femoral dentate claws, and subequal ventrites are
lines, with medial line extending to middle, sufficient to exclude Phaenocephalus from
with posterior edge deeply notched medi- allotherfamiliesofCucujoideaexceptPhal-
ally. Metacoxa large, transverse, almost acridae. With the shape ofthe tegmen and
contiguous. Metendosternite with furcal the almost completely reduced tentorium it
arms divergent at about 90 degree angle, seems that thisgenus can easily be included
anterior tendons short, curved laterally at in this family. Moreover, I have observed
apex. Leg with trochanter small, subtrian- elongatetormaefusedapicallywithelongate
gular; femurwidestatmiddle, progressively labial rods only in other phalacrids. This
longer posteriorly, fore and midfemur condition has not, to my knowledge, been
slightly longer than tibia, hind femur sub- reported previously forcucujoids. It maybe
equal in length to hind tibia; tibia elongate, related to the reduced tentorium or it may
subcylindrical, with numerous stout spines, beausefulphylogeneticcharacteratanother
especially along posterior edge, tibial spur level.
formula 1-2-2, fore spurshort, stout, barely Theonlyotherphalacrid with similartar-
longerthan tibial spines, midspurs distinct, si and tarsal claws is the Australian Phal-
unequal in length, hindspurs distinct, sub- acrinusBlackburn. SpeciesthatI examined,
equal in length; tarsi (Fig. 7) 4-4-4, all tarsi however, arelargerthatPhaenocephalus, the
320 PROCEEDINGS OFTHE ENTOMOLOGICALSOCIETY OFWASHINGTON
lllmJIINIIIIIIIIJIlMMIlMimillllll'rT
10
Figs. 1-10. Phaenocephalus castaneus. 1, Pronotum. 2, Wing. 3, Cranium, ventral. 4, Labrum, left side
ventral, right side dorsal. 5, Mandible. 6, Maxilla. 7, Tarsus. 8, Labium. 9, Tegmen. 10, Median lobe.
VOLUME 93, NUMBER 2 321
elytral punctures are more distinct, without thispieceand contributed useful comments
punctation on the intervals, and the lateral and suggestions.
margins of the elytra are distinctly and My visitsto London and Pariswere fund-
sometimesbroadlyexplanate. Itislikelythat ed by an Ernst Mayr Grant from the Mu-
these two genera are sister groups or that seum of Comparative Zoology, Harvard
they should eventually by synonymyzed. University, Cambridge, USA. These visits
Although the phylogenetic importance of would have been impossible without an ad-
some of these features is still uncertain, it ditionalgrantthattransportedmeacrossthe
is reasonable to include Phaenocephalus in Atlanticfrom theOfficeofInternational Re-
Phalacridae until a comprehensive phylo- lations, Smithsonian Institution, Washing-
genetic arrangement of this family is pro- ton, U.S.A. to study the phylogeny of the
posed. Even with such a system, the precise cerylonid series of Cucujoidea with S. A.
position ofthis genus will be tentative until Slipinski in Warsaw, Poland.
the remainder ofthe types are discovered. This paper is dedicated to the late Don
Then we may more confidently discuss the Whitehead in recognition ofhis numerous
placement of Phaenocephalus by utilizing contributions to systematic entomology.
specimens that have been compared with
Literature Cited
this material.
Acknowledgments Crowson, R. A. 1955. The Natural Classification of
the Families ofColeoptera. Lloyd, London. 187
I am deeply indebted to M. Kerley, The pp.
Natural History Museum, London, En- Lawrence, J. F. 1982. Coleoptera, pp. 482-553. In
gland, and N. Berti, Museum National Parker, S. P., ed.. Synopsis and Classification of
LivingOrganisms. Volume2. McGraw-Hill, New
d'Histoire Naturelle, Paris, France, for pro-
York.
viding material for this study. Both indi- Matthews, A. 1899. A Monograph oftheColeopter-
viduals spent longhours searching forthese ous FamiliesCorylophidaeand Sphaeriidae. Jan-
specimens in their respective institutions. I son & Son, London. 220 pp.
am grateful to H. Sasaji forgenerously pro- Paulian, R. 1950. Les Corylophidae d'Afrique. Me-
viding me with a specimen o{Phaenoceph- moires de I'lnstitute Francais d'Afrique Noire.
Number 12. 126 pp.
alus castaneus from his personal collec- Sasaji,H. 1985. Phaenocephalidae,p. 230,pi. 37, fig.
tion and to J. F. Lawrence for a gift of 2\. In Kurosawa, Y., S. Hisamatsu & H. Sasaji,
Phalacrinus. I thank S. A. Slipinski for a eds.. The Coleoptera ofJapan in Color. Volume
spiriteddiscussion ofPhaenocephaluswhile 111. Hoikusha PublishingCo., Osaka. 500 pp. [in
Japanese]
visitinghislaboratoryand forhiscomments Wollaston, T. V. 1873. On newColeoptera from Ja-
on this manuscript. R. A. Crowson, J. F. pan (part). Entomologists' Monthly Magazine 10:
Lawrence, and W. E. Steiner also reviewed 167-168 (part).