THE NORTH AMERICAN CERYLONIDAE ( COLEOPTERA : CLAVICORNIA ) By

The Cerylonidae are a amily o small to minute beetles (usually 2 ram. or less) which occur most commonly in fo.rest litter and under bark. At present, there are about 4o genera and over 3oo described species known rom all o the major zoogeographic regions. Crowso.n (I955) qrst recognized the Cerylonidae as an independent clavicorn family, including the cerylonines and murmidiines, as well as Euxestus and its allies; but these groups have been treated a.s tribes of the heteromerous amily Colydiidae by both Hetschko (I93O) and Arnett (1968). In their world generic revision of the family, Sen Gupta and Crowson (1973) added .4nommatus Wesmael, ./tbromus Reitter, and Ostomopsis Scott, while transferring Eidoreus Sharp (--Eupsilobius Casey) to the Endomychidae. The present paper consists of a revision of the IO genera and 18 species of Cerylonidae occurring in America north of Mexico. With respect to the composition of the family and that of its major subordinate groups, we have followed the classification presented by Sen Gupta and Crowson; the interrelationships among the subgroups, however, are still obscure, so we have treated the Euxestinae, Anommatinae, Metaceryloninae (not North American), Murmidiinae, Ostomopsinae, and Ceryloninae as independent subfamilies. The following abbreviations have been used in keys and descriptions" PLmpronotal length, PW--pronotal width, EL--elytral length, EWmelytral width, and TL--sum of PL and EL. The word "length" refers to the total length, including the head, and is

physiidae, and Lathridiidae.The broadly .closedmiddle coxal cavi- ties, characteristic tentorium, more or less compact antennal club rarely more than 2-segmented, and simple tarsi will distinguish the cerylonids from most o]? the above groups.Corylophids have similarly closed mesocoxal cavities, but the tentorium is reduced, the antennal club is 3-segmented and relatively loose, and the second tarsal seg- ment is oxCten lobed.Although cerylonids were ormerly included in the ]?amily Colydiidae, members of the latter group are easily dis- tinguished by having the basal three or four ventrites connate, the aedeagus o the pseudotrilobe type, and the trochanters strongly heteromeroid or the antennal insertions concealed.
Cerylonid larvae are elongate or oval in shape, without or with weakly developed frontal sutures, and with o, 2, or 3 ocelli on each side ot? head.The antennae are relatively short, with the senso.ryPsyche [June by one of us, but the particles were often impossible to identify.
The larval gut contents of the Euxestus from bat guano contained a variety of hyphal sections and spores, while numerous darklypigmented spores of a single type were tound in the adult proven- triculus of a Philothermus collected under pine bark in Mexico.The proventriculus of Ceryl,on castaneum collected on dried mushrooms from a tree was filled with sculptured basidiospores, while that of the same species taken under conit.erbark contained sections of hyphae, as well as spores.Those forms with piercing-sucking mouth- parts have been assumed to be predators (Besuchet, 1972), but as yet there is no direct evidence supporting this hypothesis (see below).
Perhaps the most interesting teature of the subfamily Ceryloninae is the occurrence of piercing-sucking mouthparts in all known larvae and in adults of over 5o species in at least six genera.This condition may have evolved twice in the larval forms and several times in the adults (see below).The most highly specialized type of piercing apparatus in adult cerylonids occurs in the genus Cautomus and has been described and illustrated by Besuchet (1972).In this group,.
the labrum-epipharynx and the labium together form an elongate, tubular beak, in which are contained four pairs of stylets, each set consisting of a mandible and its lacinia mandibularis (prostheca), a galea, and a lacinia.The molar area of the mandible is absent, while both the mandibular apex and the prostheca are long and blade-like.The maxillary stylets are extremely long and thin and are attached to a basal fulcrum which allows them to.be protracted; the lacinia bears fine recurred teeth and the galea is provided with hairs at the apex.Finally the pharynx is enlarged, forming a pharyngeal pump.This type of condition also occurs in Axiocerylon and its relatives, in the New World Cerylcautomus, and in a few species of Lapethus.
Although the mechanics have not been studied, it is obvious that this represents a piercing-sucking organ similar to those found in various Diptera and Hemiptera.Besuchet (1972) also discussed the transition from normal chew- ing mouthparts found in Cerylon to those in Cautomus through forms such as Ectomicrus, which exhibit an elongation of the labrum, mandibular apex, galea, and labium.The same type of intermediate condition may be found in species of Lapethus (figs.I-I2) and in certain Philothermus.In most of these species, the labrum and labium are somewhat elongate, the galea and lacinia are both stylet- like, the mola is usually present, and the mandibular apex retains two The larvae ot cerylonines have two different types o feeding mechanism.In Philothermus and Lapethus, the labrum and labium i:orm a tubular beak, while the mandible and the mala are.both modiiqed into long, blade-like stylets.In addition, the pharynx is enlarged and a pair o( salivary glands extend back into.the thorax i:or a distance equal to the head length.These, like the Cautomus adults, have a well-developed piercing-sucking apparatus.The larval head, however, is strongly hypognathous, and not prognathous like that ot: the adult.The second type o. eeding apparatus is found in the larvae o Cerylon.Here the head is opisthognathous with the labium short and used to the thorax, and the mandibular and max- illary stylets completely enclosed within the head and apparently attached to a heavily sclerotized internal framework.This condition is remarkably similar to that tound in the entognathous apterygote insects, such as the Diplura, Protura, and Collembola (Tuxen, 959).
As mentioned above, the actual ood source o these small and uncommon insects is otten difficult to.ascertain, and this is especially true i:or those species ingesting fluids, it: this is the case.Although it would be an obvious conclusion that those cerylonids with piercing beaks are predators on small arthropods and nematodes, it is also possible that the beaks penetrate wood or fungal hyphae or that these substances are digested extra-orally.Another possibility is that spores or other objects less that 5 or 6 microns in length are moved into the labral-labial tube by suction or by the action of the seti}erous galea; the apical openings ot those beaks examined were at least 8 microns wide.Another matter to consider is the normal position o the head.In most cf the adults with piercing beaks, the head is somewhat prognathous, so that predation would be possible on active prey species ot: various sizes.Most cerylonine larvae, however, are Psyche [June strongly hypognathous, while that o Cerylon is opisthognathous with entognathous stylets.It is less likely that either larval type would be predaceous.It is hoped that observations on living specimens will provide more inorma.tionon the eeding habits o both larvae and adults.
The evolution o piercing-sucking mouthparts has occurred at least two more times in the Coleoptera.Leiodid beetles o the genus Myrmecholeva Lea, which occur with ants in Australia have a relatively robust, piercing beak and prognathous head, and they may be predators on ant l'arvae.The maxillary stylets are well-developed and blade-like in this group, but the mandibles are much shorter than the beak and do not appear to unction as piercing organs.In the :amily Eucinetidae, members o an undescribed genus rom the New World are extremely small (less than ram.) with a hypognathous head and a labral-labial beak with an opening o: about 5 microns; the mandibles in these insects are even urther reduced, but the max- illary stylets are very long and ine.A detailed comparison o pierc- ing-sucking eeding mechanisms in beetles will be included in a paper now in preparation by J. F. Lawrence and T. F. Hlavac.
Peck, and W. Suter.We are grateful to R. Crowson, J. Doyen, T. Hlavac, A. Newton and R. Silberglied or advice given during portions of this study.We also wish to thank Virginia.Phear, Marilyn Pearce, Jennifer Slade, and Margaret Thayer for typing and proofing, and to F. 3/I.Carpenter for a critical reading of the manuscript.
Key to the Genera of North American Cerylonidae I. Frontoclypeal suture present; terminal segment o( maxillary palp at least 2 X as long as and more than 0.5 X as wide as segment 3, which is shorter than or subequal to segment 2; last ventrite not or very finely crenulate 2 Frontoclypeal suture absent; terminal segment o maxillary palp aciculate, shorter than and less than 0.5 X as wide as segment 3, which is longer than segment Lateral margins of pronotum crenulate; eyes absent; body elon- gate, TL/EW about .5;elytra with 7 rows of punctures bear- ing very short, fine hairs; procoxal cavities very narrowly sep- arated, the intercoxal process depressed and less than o.2 ( a coxal width

Philothermus Aub6
Procoxal cavities broadly closed behind, the postcoxal bridge more than 0.5 X as wide as cavity (ig.o) intercoxal process of prosternum strongly widened posteriorly; mesosternum flat or slightly convex; antennae o-segmented with I-segmented club; lateral margins of pronotum not visible for their entire lengths from above Cerylon Latreille Euxestinae Grouvell.e As used here, this group is equivalent to the tribe Euxestini de- scribed and delimited by Sen Gupta and Crowson (I973), and in- cluding the Cycloxenini and Tachyoryctidiini o Jeannel and Paulian (I945).Sen Gupta and Crowson admitted that those characters shared by the Euxestini, the Anommatini, and the Metacerylonini o Heinze (I944) are, or the most part, primitive or the family, and that all three tribes might be given subfamily rank.Of the eight known genera, only one, Hypodacne, occurs in America north o Mexico.The genus Euxestus Wollaston, however, includes a widely distributed Neotropical species which might be expected to occur in southern Florida (see below).
Sen Gupta and Crowson resurrected LeConte's name for a group of species, usually included in E,uxestus.which have simple tarsi, lacking the characteristic ventral lobe on the first tarsal segment.
In addition to the North American H. punctata, several Australian and New Zealand species are included.
Hypodacne punctata LeConte (Fig. 2o) This species is easily distinguished from other North American cerylonids by the highly convex and oval body, which is longer than 1.5 mm., widely separated and posteriorly closed procoxal cavities, distinct frontoclypeal suture and characteristic antennal club, .and the lack of pronotal antennal cavities, femoral lines on the abdomen, and crenulations on the last visible ventrite.The related Euxestus erithacus Chevrolat occurs in the Greater Antilles and might be found in southern Florida, but that species is smaller in size and bears a distinct ventral lobe on the first tarsal segment, a feature absent in Hypodacne.
Distribution.Known from scattered localities throughout eastern North America, from southern Ontario to Florida and west to Kansas and Texas.
Biology.According to Stephan (968), this species occurs in the galleries of carpenter ants (Camponotus) in southern Ontario.Speci- mens were observed crawling on walls of carpenter ant galleries in oak, elm, and beech.The larva of H. punctata is unknown, but that of the Australian species, H. bivulneratus (Lea), was described by Sen Gupta and Crowson (I973: 38I) from specimens collected in leaf litter and decayed wood.

Anommatinae Ganglbauer
This subfamily includes the two Palaearctic genera A bromus Reitter and A nommatus Wesmael, the latter o which has been introduced into North America.Ganglbauer (899) originally con- sidered this tribe to be related to cerylonids, but Crowson (955) Psyche [June placed the group in his Merophysiidae.Sen Gupta and Crowson (I973) recognized its affinities to the Cerylonidae, and especially the Euxestinae, ater the description of the Anommatus larva by Dajoz 968).
This genus is native to Europe, where it is represented by 49 speci:es.It has been recently revised by Dajoz (965).
Distribution.Fairly widely distributed in Europe and extending at least to Turkey and Algeria.Recorded also from Madeira, St.
Biology.Adults have been collected rom litter and damp soil under railroad ties, under stones, around grass roots, and from grass cuttings.The Illinois specimen was taken in a tree hole.Larvae have been found in soil at the base of a tree.and in leaf mold (Dajoz,   968; Sen Gupta and Crowson, 973).No males have been ound in North America, and Cooper (962) and Peck (I972) have postulat.edthat our populations may be parthenogenetic.

Murmidiinae Jacquelin DuVal
This corresponds to the Murmidiinae o Sen C-upta and Crowson, except for the exclusion o.f Ostomopsis (see Ostomopsinae below).
The group may be characterized by the presence o a rontoclypeal suture and prothoracic antennal cavities, and it includes the three genera Murmidius, Mychocerus, and Botrodus, all of which are represented in North America.Casey (895) also included the Lapethini and the Eupsilobiini, with the single genus Eupsilobius.
According to Sen Gupta and Crowson ( 973), the lapethines should be transferred to the Ceryloninae, whil.e Eupsilobius Casey, a junior synonym o( Eidoreus Sharp, should constitute a distinct subfamily o Endomychidae.
This genus contains ive Neotropical species and five from the Old World.One ,of the latter, M. ovalis, has been introduced in stored products to many parts of the world, including North America.Hinton (942b) provided a key to the Old World species.
Biology.Usually associated with stored products and often found in granaries and warehouses.Reported from ruits and seeds, Indian corn, rice, wheat, flour, hay, straw, gall-nuts, oak-galls, ginger, .andpepper, and also collected in dead leaves and cut grass (Hinton, 945 Halstead, 968).The larva, which is flat and onisciform, with barbed setae, has been illustrated by Boving and Craighead 93 and Halstead 968), while the last author figured the pupal cocoon and gave additional information on life history and develop- ment.

Mychocerus Erichson
Mychocerus Erichson, 1845" 292.Type species, by monotypy, M. [errugineus Erichson.This New World genus contains five species, two.o which occur in America north o Mexico.The described Seychelles species, M. alluaudi Grouvelle, was transferred to Lapethus by Hinton (936).Grouvelle (t895) also included Philothermus gibbul.usReitter (t876), without giving his reasons; although Reitter's type has not been seen, the description o.f gibbulus could hardly apply to a species of Mychocerus.The species o Mychocerus may be dis- tinguished by the small size and flattened .orm,9-segmented antennae with a t-segmented club, anteroventral antennal cavities, and seria.teelytral punctation.The literature is somewhat conused as to the type species for this genus.In the original description, Erichson mentioned that three species were included in his genus, but he named one from Cuba., "'M.[errugineus Nob."Since the genus was described in the same paragraph, this must constitute the species description as well, and thus M. ferrugineus is the only included species and the type.We have not seen the Cuban species, but we are assuming that it is congeneric with M. depress,us, which LeConte (t869) placed in Mychocerus, and which is usually considered the type.

Key to the North American Mychocerus
Sides of pronotum gradually converging near apex, not angu- late; lateral lines o mesosternum subparallel or most o their lengths, abruptly converging anteriorly, each line posteriorly forming an abrupt angle (about 2o) with the metasternal line (rig.2); apical edge of prosternum slightly emarginate; size smaller, le.ngth usually less than ram.eastern North America.

Myehoeerus arizonensis, New Species
This species may be distinguished from either M. depressus or M. pili[er Lewis (888: 24) by the larger size and by the shape o: the pronotum, which is more abruptly narrowed apically.From M. depressus it also differs in the form o the mesosternal lines.Description.Length .-.3 mm.Color reddish-brown; vestiture o: short, fine, decumbent, yellow ha.irs, which .are barely visible under 25 X magnification.Antennae with segments 4-6 distinctly serrate, the club about 1.35 M as long as wide.Pronotum about 0.48 >( as long as wide, sides subparallel for basal three-fourths, abruptly converging apically, the apical angles produced a.nd subacute; lateral margins distinct and slightly elevated, especially towards base; disc slightly convex; punctation somewhat irregular, coarser than in M. depressus.Elytra about I.I5 >( as long as wide and 3.o )< as long as pronotum, widest at apical fourth; rows of punctures lightly im- pressed and not distinct, the punctures smaller than those of M. depressus, intervals shiny, but with a fine reticulation.Apical edge of prosternum distinctly concave; mesosternal lateral lines gradually converging anteriorly, each line posteriorly forming an angle of about I5 o with the line of the metasternum (fig.3).Aedeagus about o.6 X as lo,ng as abdomen, parameres longer and narrower than those of M. depressus, about 3 ( as long as wide at base (figs.5-6).
Distribution.Southeastern Arizona.and southern Sonora, Mexico.In Arizona, known from low elevations in the extreme southern pa.rt of the state and from higher elevations in the Chiricahua Mountains.Biology.Individuals from the lowland Arizona, population were taken under dry mesquite bark and were not associated with running wounds.Those from the montane locality were collected under oak ba.rk.
Mychocer.usdelressus (LeConte) (Fig. 22) Murmidius delressus LeConte, 1866" 376.Type locality: Washington, D.C. Holotype, No. 6829 M.C.Z.This is the smallest cerylonid in our fauna, seldom exceeding ram. in length.It is very similar to M. arizonensis, but may be distin- guished by the smaller size, more gradually na.rrovced pronotal apex, subparallel mesosternal lines which form a more abrupt angle with the lines on the metasternum, and shorter, broader parameres (about twice as long as wide at base).The pronotal punctation in M. delressus is/]ner than that of arizonensis, while the elytral punctures are larger and more distinct.According to.Lewis (I888), the Gua- temalan species, M. pilifer, is about the same size, but has a differ- ently shaped pronotum and less distinct elytral punctation.
Distribution.Known from scattered localities in eastern North America, from southern Ontario south to.Florida and west to eastern Texas.
Biology.Adults have been collected under ungus-inested bark of oak, and also under bark o.f beech and hickory.The larva is LlnkI1oWTI.
This species resembles a small lapethine with respect to its oval and highly convex body, conused elytral punctation, and ventral, rather than dorsal antennal cavities.Its relationship to.Mychocerus and Murmidius is indicated by the presence of a fr.ontoclypeal su- ture, murmidiine mouthparts (galea and lacinia not elongate, terminal segment o maxillary palp not aciculate), and Mychocerus-like aedea- gus.The species may be distinguished rom the other murmidiines by the ant'eriorly rounded mesosternum, 8-segmented antennae, and lack o metasternal suture.
Distribution.Known only rom Florida and Texas.
Biology.H/bitat and immature stages unknown.This monotypic subfamily is considered to be a tribe of Murmi-  diinae by Sen Gupta and Crowson (1973), but its relationship to the latter group is somewhat unclear.It is here treated as an inde- pendent subfamily.
Ostomol)sis Scott This genus was described by Scott (I922: 250-253, pl.I9, fig. for the Seychelles species 0. solitaria, and has been redescribed and illustrated by Sen Gupta and Crowson (1973: 4oo-4o2, figs.[43][44][45][46][47][48][49] on the basis of new material from New Caledonia and Florida.There is little to add to the generic description provided by the latter authors, except that the hind edge of the pronotum has a raised margin and the scutellum is transverse.Also the antennal club is broader than their illustration suggests, and it appears to be pinched in the middle due to the presence of lateral sensillae (see fig. 4).
Species of Ost, omopsis may be distinguished from other Cerylonidae by the frontoclypeal suture, internally and externally open pro- coxal cavities, and 3-segmented tarsi, and the lack of femoral lines, antennal cavities, and a transverse line on the vertex.Ostomopsis solitaria Scott is known only from the Seychelles, but specimens ap- pearing to be conspecific have been seen from New Caledonia and the Loyalty Islands.The Neotropical species described below extends the range of the genus into the New World.Ostomopsis neotropicalis, New Species (Fig. 24) This species may be distinguished from O. solitaria by the smaller size (less than I.IO mm.), somewhat more elongate body, and nar- rower, less explanate, pronotal margins with more irregularly serrate edges.
Description.Length o.95-1.1omm.Color yellowish-brown; vesti- ture of dense, suberect, yellowish hairs, which are about three-fifths as long as scutellar base, intermixed, especially at sides, with much longer hairs exceeding scutellar base in length.Antennal club (fig.4) about as long as wide and twice as wide as the segment preceding it.Pronotum about o.6o X as long as wide, widest at posterior third, sides weakly rounded, apical angles slightly produced and acute; lateral margins moderately broad and explanate, but less so than in O. solitaria, the edges coarsely and irregularly serrate; disc slightly 1975] Lawrence & 8tephan--Cerylonidae I47 convex, slightly flattened mesially, punctation moderately coarse and dense, the interstices shiny.Scutellum transverse.Elytra about 1.42 )< as long as wide and 2.50 X as long as pronotum; sides weakly rounded, almost parallel basally, the margins somewhat explanate, with nely, but sparsely and irregularly serr.ateedges; punctures o two types: large, somewhat rectangular ones, closely packed in regu- lar rows, and very ne ones occurring in the intervals, both types bearing suber.ecthairs.Aedeagus about as long as abdomen; tegmen (i]g.7) with fused parameres and very long median strut; median lobe about as long as tegmen, very narrow, with long basal struts.
Biology.Possibly associated with rotten palm wood.Specimens o O. solitaria rom New Caledonia and the Loyalty Islands were collected under bark.

Ceryloninae Billberg
This subfamily includes all of the remaining cerylonids, which are characterized by the lack o a rontoclypeal suture and the pres- ence of relatively long and slender maxillary lobes, aciculate maxil- lary palps, and a distinctly crenulate margin on the last ventrite.
The last feature, which may be present but much less obvious in cer- tain murmidiines, is actually a series o vertical ridges on the in- flexed margin o.f the ventrite (fig. ); these form an interlocking mechanism with a correspo.ndingseries beneath the elytral apices.
All known cerylonine larvae and adults in at least six genera have evolved highly modified, piercing mouthparts, with an elongate, pointed labrum, and mandibular and maxillary stylets (see p. I34)- and the presence of lateral antennal cavities on the anteroventral portion o1: the pronotum; three genera are included at present, but two ot: these are synonymized below.The remaining genus Laethus is closely related to certain Cerylonini, and the larvae are more similar to those o1: Philothermus than the latter are to the larvae o1: Cerylon.The Aculagnathini are a diverse group which have one eature in common-the possession o1: piercing mouthparts.In the Indo-Aus- tralian genus Cautomus and its relatives, the prosternum is raised in the middle, so that a shallow antennal cavity is 1:ormed on either side, and the dorsal sur1:ace o the body is relatively simple.The Asian and A1:rican dxiocerylon, Dolosus, and Thyroderus, on the other hand, are characterized by having the prosternum produced orward to form a chin-piece, which fits against the clypeus to con- ceal the mouthparts, and by the possession o1: deep antennal cavities, dytral ridges, and pronotal grooves.In both o1: the above groups, the labrum tapers to a sharp point, and a tube is ormed to contain the mandibular and maxillary stylets (see p. 134).In the New World, Cerylcautomus ltoridensis resembles the Cautomus group in having shallow median antennal cavities and an acute labrum, but this species differs in the 1-segmented (instead o 2-segmented)   antennal club.In Central America and the West Indies, a number o( undescribed orms have been seen which resemble C. ltoridensis in most respects, but which vary considerably in the development o1: those characters considered to be diagnostic o1: the Cautomus group or ot: aculagnathines in general.In one case, the ant.ennal cavities are absent and the labrum is elongate but not acute or tube-like, while in another the antennal cavities are well-developed but the mouthparts are not at all o1: the piercing type.Finally, the South American Glyltololus resembles a large member of the Axiocerylon group, but thee characteristic chin-piece and antennal cavities are absent, the antennae are 3-segmented, and the labrum is not .acute.The great variation within this tribe, the existence o.1: a number o1: (orms with int.ermediate types o1: mouthparts, both within the tribe and within cerylonine genera, such as Ectomicrus, and the occurrence o1: piercing beaks in the lapethines and in cerylonine larvae, all sug- gest that the Aculagnathini may be polyphyl.eticand that the diag- nostic piercing type o mouthparts may have arisen independently in A more detailed analysis o the group and the related Cerylonini, based on characters unrelated to the feeding apparatus, may shed more light on the problem.The Cerylonini include the remaining dozen or so cerylonine genera which have not been placed in the above tribes, and these also orm a diverse assemblage.The large genera Cerylon and Philo-  thermus are discussed below, while the smaller exotic groups are covered in Sen Gupta and Crowson (I973), Hinton (I942a), and Heinze (1944).
There has been considerable confusion concerning the constitution of this genus, and both Heinze (944) and Sen Gupta and Crowson (973) have attempted to clarify its limits and to.distinguish the included species from those of Cerylon.The latter authors have defined the group primarily on the basis of prosternal characters and have included a number of forms with o-segmented antennae and a I-segmented club.Among these are members of the genus Ecto- micrus Sharp (1885) and a group .ofAfrican Cerylon-like forms.
Although Ectomicrus was synonymized with Philothermus by Reitter (89o), Heinze (944) ound several characters to distinguish it, in addition to the I-segmented antennal club.The African torms mentioned (at least P. camerunense Sen Gupta and Crowson) have well-developed femoral lines on the abdomen and appear to have more in common with Philothermopsis than Philothermus, in spite of tarsal differences.As presently conceived, Philothermus includes about 5o species, which are characterized by having a more or less elongate form, 4-segmented tarsi, no prothoracic antennal cavities, procoxal cavities open or narrowly closed behind, and the prosternal process not or only slightly widened and more or less rounded at apex.In addition, most of the species have II-segmented antennae with a 2-segmented club and no femoral lines on the first ventrite.
In North America, the genus is represented by a California and a Florida endemic and a widespread eastern species.
Key to the North American Philothermus I. Body more elongate, PL/PW more than 0.82 and EL/EW more than 1.58; each elytron with 7 rows of punctures (not 2. counting that along lateral edge); lateral margins of pronotum narrow; procoxal cavities narrowly closed behind; California.occidentalis, n. sp.
Body shorter and broader, PL/PW less than o.82 and EL/EW less than 1.58; each elytron with 8 rows of punctures; lateral margins of pronotum broad and explanate; procoxal cavities open behind; eastern North America 2 Elytral vestiture consisting of numerous erect hairs, which are usually much longer than the diameter of an elytral puncture; pronotal punctation coarser and denser, the punctures usually separated by less than the diameter of one; lateral margins of pronotum distinct crenulate and lateral margins of elytra visible from above almost to apex; Florida puberulus Schwarz Elytral vestiture consisting of very short hairs, which are shorter than a puncture diameter, and a few scattered longer hairs; pronotal punctation finer and sparser, the punctures usually sep- arated by more than the diameter of one; lateral margins of pronotum relatively smooth and lateral margins of elytra visible from above only for .anteriorhalf glabriculus LeConte Philothermus glabriculus LeConte (Fig. 25 Philothermus glal,riculus LeConte, 1863: 69.Type locality: Pennsylvania. Leetotype, 6, No. 6827 M.C.Z.(selected by J. F. Lawrence, April,   1975).This species may be distinguished from the Floridian P. luberulus by the larger size (I.9-2.3 ram.) and somewhat broader form, shorter elytral hairs, finer and sparser pronotal punctation, and relatively smooth and even lateral prono.talmargins.The western P. occi- dentalis is more elongate, with narrower pronotal margins and closed procoxal cavities.This is probably the most common cerylonid in eastern North America and it may occur with Cerylon unicolor, which is easily distinguished by the much narrower form and I-seg- mented antennal club.
Distribution.Widespread throughout eastern North America from southern Ontario and Maine, south to northern Florida .andwest to Oklahoma and Texas.
Biology.Adults have been taken under the bark of ricer, Carya, Fagus, Pinus, Quercus, Tsuga, and Ulmus.Also taken under boards, in rotten logs, sawdust piles, malt traps, leaf litter, tree holes, and forest debris.This species is characterized by the elongate body, narrow pronotal margins, 7 rows of elytral punctures, narrowly closed procoxal cavi- ties, slightly expanded pronotal process, deeply concave mesosternum, and elongate metasternum.
Description.Length 1.8-2.2ram.Color reddish-brown; vestiture o short, erect hairs, which are visible under 25>( magnification. Pronotum about o.85 X as long as wide, widest at basal fourth, sides slightly converging to rounded apical angles; disc moderately convex, lateral margins narrow and somewhat irregular, the edge slightly crenulate at base; punctation airly coarse and dense, punc- tures near center about o.2 X as large as scutellar base and usually separated by less than a diameter; interstices smooth and shiny, with scattered very ine punctures.Elytra about 1.67 > as long as wide and 2.25 X as long as pronotum, widest at anterior third, each elytron with 7 rows o punctures (not counting marginal row), the punctures .aboutas large as those on pronotum, deeply impressed, and fairly closely and irregularly spaced, intervals slightly convex, smooth and shiny, with scattered, very ine punctures.Prosternal process slightly na.rrovcer than a coxal diameter, very slightly ex- panded posteriorly and broadly rounded at apex; coxal cavities nar- rowly closed behind (ig.9).Mesosternal concavity fairly deep, without a median carina, the process truncate at apex.Metasternum slightly longer than first ventrite.Aedeagus (fig.8) with long, narrow parameres; median lobe about o.75 X as long as abdomen.
Ty#es.Holotype, Mather, Tuolumne Co., CALIFORNIA, June Distribution.Mountains of northern California, and south in the Sierra Nevada at least to Mariposa 'County.
Biology.The Mather series was sifted from conifer debris to- gether with Cerylon californicum.
Philothermus occidentalis is not very closely related to its eastern congeners, as evidenced by the pro.sternal and mesosternal characters, as by the elongate parameres, which are extremely reduced in both puberulus and glabriculus.
This species is easily distinguished from P. glabriculus by the vestiture of numerous, long, erect hairs and the distinctly crenulate lateral pronotal ma.rgins.It is also smaller in size (r.6-.9 mm.) and somewhat more elongate, with coarser and denser pronotal punc- tation.
Distribution.Georgia, Florida, and the Greater Antilles.Re- ported by Hinton (936) from Mexico, Guatemala, and Panama.. Biology.Adults collected under bark of Pinus and in a berlese sample from a swamp floor.
The Neotr.opical distribution o.f this species must remain tentative until a complete revision is undertaken, since there appears to be a complex of tropical species related to P. puberulus.Specimens from Cuba, Puerto Rico, and Hispaniola appear to be conspecific with the Floridian population, but most of the specimens we have examined from Mexico and Central America are not the same species.Horn (878) also reported this species from Arizona, but we have seen no specimens to verify this record.

Cerylcautomus Sen Gupta and Crowson
Cerylcautomus Sen Gupta and Crowson, 1973: 439-441.Type species, by original designation, C. itoridensis Sen Gupta and Crowson.This ffenus was based on a single Floridian species and several unnamed forms from Jamaica, Hispaniola, Puerto Rico., Trinidad, and British Guiana.All members of the genus have at least vaguely defined antennal cavities, similar to those in Cautom.us,located later- ally on the prosternum (not on the hypomera) and defining a median, raised, prosternal plate.In addition, the procoxal cavities are open Psyche [June behind, the antennae are o-segmented with an elongate, I-segmented club, and the labrum is elongate and narrowed, or sometimes acute, at the apex.
This species is easily distinguished from other North American cerylonines by the small size (less than .25 ram.) and elongate i:orm, o-segmented antennae with a -segmented club, prosternal antennal cavities, and acute labrum.
Distribution.Known only rom northern Florida.Biology.Recorded rom an old sawdust pile (more than 2o years old) at the edge o a cypress swamp.Other members of the genus have been extracted with Berlese funnels from forest litter and rotten logs.
Lapecautomus Sen Gupta and Crowson, 1973" 409.Type species, by ori- ginal designation, L. dybasi Sen Gupta and Crowson.New Synonymy.line.Likewise, the condition represented in the three described species of Lalecautomus piercing bea.k with mandibular and max- illary stylets----is on.e which occurs throughout the subfamily in both larvae ,and adults, so that a gr.oup, .evenat the generic level which is defined on the basis of this feature alone, may well be polyphyletic.
Within the genus Lalethus (in the broad sense), there is a general trend towards the elonga.tion of the labrum, and in most forms examined it is somewhat elongate and deeply emarginate at the apex, as in L. discretus (fig.I:).In Lal)ethus (-Lal)ecautomus) striatus, th.e labrum is even more elongate than in discretus, but it is still emarginate at the tip (fig.II).In L. dybasi, however, the apex is pointed.A similar trend may be found in the mandibles and max- illae.With respect t.o other characters not involved in th.e piercing apparatus, the three species presently included in Laecautomus do not appear to be closely related and ma.y well be derived from two or more stocks of typical Lal)ethus.Lapecautomus mexicanus, for instance, is a globose, highly convex form, which is almost impunc- tare and glabrous, and which has the type of pr.o-and mesosternum characteristic of a Lytol)eplus.In co,ntrast, L. striat.us is more .elon-gate and flattened, with distinctly seriate elytral punctation, erect vestiture, and a raised lateral pronotal margin; in all these respects it resembles the type of Lal)ethus, L. discretus.For these reasons, we recognize the single genus Lapethus, and we suggest that the evolution o.f compact, globose f.orms and forms with piercing mouthparts represent two independent trends occurring in more than one lineage.
The genus Lapethus, as h.ere delimited, contains about :5 described species, and a number of undescribed forms have been seen, especially in the New World.The group is primarily tropical, and the North American species are restricted to humid areas along the Pacific coast and in the mountains of the Southeast.
Key to the North American Lal)ethus Pronotal punctation very fine and sparse, the punctures usually separated by at least two diameters; eyes reduced consisting of about O t:acets; labrum slightly longer than broad at base; size smaller, TL less than 1.75 mm.; Great Smoky Mountains and vicinity striatus (Sen Gupta and ,Crowson) This species is .characterizedby the oval, somewhat flattened orm, reddish color, distinctly seriate elytral punctation, fine, ,erect hairs easily visible under o)< magnification, distinct lateral pronotal mar- gins, 8-segmented anlzennae with a I-segmented club, .andlarge antero- lateral antennal cavities on the underside of the pronotum.The.pro- coxal cavities are fairly narrowly separated and distinctly open behind, while the prosternal pro.cess is parallel-sided .andstrongly rounded at apex.Philothermus glabriculus is superficially similar but more elongate and parallel sided, with different antennae and no antennal avities.The species may be distinguished Cro.m the related L. striatus by the larger size (2.o-2.3 mm.), coarser and denser pronotal punc- ration, shorter labrum, and fully developed eyes and wings.
Distribution.P, aciflc Coast, from western Washington (King County) to central co,astal California (Santa Cruz County) and the Sierra Nevada (Tuolumne and E1 Dorado Counties).
Biology.Coll,ected in the Sierra Nevada by sifting rotten coni,f,er logs in the red rot stage.
Lapethus striatus (Sen Gupta and Crowson), New Combination Lapecautomus striatus Sen Gupta and Crowson, 1973: 412, fig. 178.Type locality: Below Ramsay Cascades, 3500', Great Smoky Mrs. National Park, Sevier Co., Tennessee.Holotype, Field Museum.This species is smaller than L. discretus (.5-.7 mm.), with the pronotum more finely and sparsely punctate, the eyes reduced to about o acets, and the wings completely absent.The labrum is also longer and narrower at the apex, and the mouthparts are more highly modified for piercing.
Distribution.Known only rom the cove forests of the Great Smoky Mountains, Tennessee, and the nearby Joyce Kilmer Forest, North Carolina.Biology.Adults have been collected in samples of leaf litter under Rhododendron and in mixed hardwood and Tsuga orest.
Distribution.Northern and eastern North America, from south- eastern British Columbia and central Alberta to southern Quebec and Maine, and south to Texas and the Carolinas.More commonly collected in the northern part of the range.
The name Cerylon castaneus has usually been associated with the species called C. unicolor in this paper, and in most collectio,ns these two names are confused, Casey's clypeale and sticticum being ap- plied to specimens with expanded pronotum and notched clypeus.
In Say's description, the point is made that the thorax is widely and slightly indented at base, .eachside o,f middl.e.This could refer to the basal narrowing of the pronotum, but it may also, refer to discal impressions.Specimens under the name castaneum in the LeCont.'ec.ollection are definitely of the type with apically expanded pronotum and notched clypeus in the male, and we are considering this as Say's species.The first specimen in the series has an orange disc (referring to southern states) and it is a female.Two other speci- mens have i:aded discs, so that the locality is uncertain.A ourth specimen bears the label "Can." and is a male.Since the species is much more .commonin the northern p,a.rt of the continent, we ha.ve chosen this last specimen as Neotype.
Cerylon eonditum, New Species This species is in the size range of C. distans, but differs from that species by the more elongate pronotum, more approximate meso- coxae, and notched clypes in the male.Cerylon unicolor is usually smaller, and less elongate, without the no.tched clypeus in the male, while C. castaneum differs in having the pronotum strongly expanded apically and the elytral intervals flat.Description.Length 2.2-2.7 ram.Color dark reddish-brown; vestiture ot very short, erect hairs, barely visible under 5oX mag- nification.Antennae about o.75 X as long as pronotal width; seg- ment 3 .25>( as long as 4; club .3X as long as wide.Clypeus slightly emarginate in 9, deeply notched in d'.Pronotum (fig.6) o.95 ( as long as wide, sides subparallel or slightly diverging, for basal three-ourths, .convergingapically to rounded angles; disc very slightly convex at middle, slightly sloping anterolaterally, with 2 Cerylon distans, New Species Fig. a9) The widely separated me.socoxal cavities will separate this species from all others in North America.In addition, the pronotum is much shorter and broader than in C. conditum, which also differs in having the clypeus notched in the male.The general form is similar to C. unicolor, in which the mesocoxae are more approximate, the size smaller, and the pronota!impressions shallower.
Description.Length 2.4-3.oram.Color dark reddish-brown to black; vestiture of very short, erect hairs, not visible under magnification.Antennae relatively short, about o.6 X as long as pronotal width; segment 3 1.4 as long as 4; club only slightly longer than wide.Clypeus slightly to moderately emarginate, but never notched.Pronotum about o.87 X as long as wide, sides sub- parallel for basal three-fourths, converging apically to rounded angles; disc moderately convex at middle, steeply sloping anterolaterally, with 2 broad and moderately deep basal impressions; punctation variable, coarser at base than apex, punctures near center about o.2 X as long as scutellar base and usually separated by less than a diameter; interstitial microsculpture very lightly impressed or absent, with scattered fine punctures.Scutellum transversely rugulose.Elytra about 1.6 X as long as wide and 2.35 X as long as prono.tum,widest at middle; striae deeply impressed, their punctures elongate and subconfluent; intervals moderately convex, shiny, and finely punctate.Prosternal process slightly concave, about 1.8 )< as wide as a coxal diameter.Mesosterna.1 process 2 X as wide as a coxal diameter, truncate at apex.Metasternum .35X as long as first visible abdominal sternite.Aedeagus (fig.I4) about 0.6 X as long as abdomen.
in the male, and the less flattened orm.Cerylon distans is larger, with more widely separated mesocoxae, while C. conditum is usually larger and more elongate, with a notched clypeus in the male..In California, this species is easily distinguished from C. californicum by the much smaller size, less distinct vestiture, and less robust orm.
Cerylon unicolor is the only species in which there appears to be pronounced sexual dimorphism in prothoracic shape, the sides in the female being parallel basally and convergent apically, while those in the male are slightly divergent .at the b,a.se ((igs.8-9).
Distribution.Northern and eastern North America and th,e Pacific Coast.Extending from southern British Columbia, south through Washington, Oregon, Idaho, and California as ar as Los Angeles County on the coast and Fresno County in the Sierra Nevada; east across southern 'Canada and the northern United States to southern Ontario and Maine; south on the Atlantic Coast to northern Florida, and west to the ooth meridian.
2; apex of last ventrite always strongly crenulate (qg.7 2. Procoxal cavities closed behind; length at least 2 ram.body oval and highly convex, with smooth, even, lateral edges and no (emoral lines; tarsi 3-segmented; body more elongate, TL/EW these genera(Cerylon, Philothermus, Lalethus)are ar rom being understood.Sen Gupta and Crowson (1973) divide the group into three tribes" Lapethini, Cerylonlni, and Aculagnathini.Members o the first group are distinguished by the oval or globose 1:orm Sharp originally placed L. compactus in a separate genus because o the widely separated pro-and mesocoxae and the coadapta.tionox the pro-and mesosternum.Hinton (1936) pointed out that the differences were only a matter o degree and that certain Lapethus, such as L. sharpi Champion, displayed an intermediate condition; on this basis he synonymized Lytopeplus with Lapethus.Sen Gupta and Crowson, however, recently resurrected Lytopeplus, stating that the only difference between species of the genus and those o Lapeth,us is the slight closure o the procoxal cavities in the o.rmer.Ater exam- ining a number o. lapethines xro.m Mexico and Central America, we are o the opinion that the eatures diagnosing Lytopeplus represent a general trend towards compaction throughout the group, and that these compact orms are not necessarily members o a single phyletic