job contact Login 

changes in the classification of the Ciidae

By John F. Lawrence
No. 5 — Comments on Some Recent Changes in the Classification of the Ciidae (Coleoptera)
Musum of Comparative Zoology, Harvard University, Cambridge, Mass.

 The family Ciidae includes at present 40 genera and about 550 species. The group has been badly neglected in the past, and although a few excellent local monographs have appeared,
the last treatment of the world fauna was published in 1848.
The European fauna has probably been more extensively studied than that of any other area, but most of the workers have not been specialists and have tended to be provincial in their approach, completely ignoring, for instance, the contributions to the Holarctic fauna made by Japanese and Americans. Lohse (1964) produced a short paper in which some of the European generic concepts were reexamined and 3 new generic group names proposed. This recent contribution is certainly welcome, since it represents the first step in clarifying the relationships within this difficult group. In the present discussion, I will review Lohse ‘s proposals and reevaluate his concepts in the light of my own work on the North American Ciidae. Since the histories of the various European generic names have been discussed previously, it may seem redundant to include them here ; I think thisis justified, however, considering that even Lohse overlooked or misinterpreted some of these historical facts.
Although the genera Entypus and Rhopalodontus representdistinct taxa whose species are only distantly related, the nomenclatural histories of the two names are so interwoven that it seems practical to discuss them together. Ropalodontus Mellie (justifiably emended to Rhopalodontus by Gaubil, 1849) was first proposed in 1847 for the species Cis perfomtus Gyllenhal, the main diagnostic character being the expanded, rounded, and externally spinulose protibial apices. In the same year, Redtenbacher described Entypus, which included only the species Cis affinis Gyllenhal (misidentified as Apate fronticornis Panzer),characterized by having 9-segmented antennae, with the 3rd segment longer than the following 3 together, and spinulose protibialapices. In the following year, Mellie included C. affinis in his genus Ennearthron, which had been described some months prior to the publication of Redtenbacher’s work, and placed Entypus
in synonymy. Mellie included the true A. fronticornis in the genus Cis, primarily because of its 10-segmented antennae.
Thomson (1863) considered Ehopalodontus and Orophius Redtenbacher (=Octotemnus Mellie) to be so different from other members of the family that he included them in a separate tribe, the Orophiina. Being a careful worker, he found several other characters which consistently distinguished the group, such as the conical, projecting, and subcontiguous procoxae, short prosternum, deep antennal sulci, and subequal 3rd and 4th antennal segments. In the same paper, Thomson noted the close similarity
between Ennearthron affine (Gyllenhal) and Cis fronticornis (Panzer), and in spite of the difference in antennal segmentation, he placed them both in Entypus. Mellie ‘s name Ennearthron could not be applied, because a very different species, Cis cornutns Gyllenhal, had been designated as its type by Demarest (1860). Kiesenwetter (1877) included Entypus as a distinct subgenus of Cis and added to it a third species, Ennearthron ivagai Wankowicz, which had been described as having spinulose protibial apices. He considered the Orophiina to be a separate family, the Orophyidae, to which he added Xylographus Mellie.
A more extreme view was taken by Seidlitz (1872), who placed these genera in a tribe of the family Anobiidae. In spite of the efforts of Thomson and Kiesenwetter, most European entomologists chose to ignore these two concepts and to rearrange the above species according to more superficial characters, such as the number and relative lengths of antennal segments. Thus, Jacquelin Du Val (1861) placed C fronticornis in the genus Rhopalodontus in spite of its totally different procoxal structure, and returned affinis to Ennearthron, because of its 9-segmented antennae. In 1915, Peyerimhoff placed Cis hicornis Mellie in Rhopalodontus on the basis of antennal and protibial characters only, and Nobuchi (1960a) described 2 more species, R. japonicus and R. tokunagai, which also appear to be in the fronticornis group. Until quite recently, then, the genus Rhopalodontus contained two distinct and distantly related
groups of species : those resembling Ennearthron affine and belonging properly to Entypus of Redtenbacher and Thomson, and those resembling R. perforatus.
The situation was clarified by Lohse (1964), who had restudied the European species carefully enough to recognize the true

relationships. By reviving the concept of Redtenbacher and Tliomson and uniting E. affinc, E.fronticornis, and R. hicornis into a single genus, he not only called attention to the apparent affinities of these three species, but also eliminated discordant elements from both Ennearthron and Rhopalodontus. In addition to the above, Lohse added Cis bidentulus Rosenhauer to the group. Because of the variation in antennal segments, he proposed the subgenus Entypocis (with C. bidentulus as its type) for those species with 10-segmented antennae, leaving Entypus affinis the only member of the nominate subgenus. Ennearthron wagai Wankowicz (1869) (mis-cited in Lohse as E. wagai Wanka) was placed in a new genus, Wagaicis, because of its very narrow and laminate prosternal process.
I agree with Lohse ‘s revival of Entypus and his inclusion of the 4 European species. In my own studies of North American Ciidae, I have encountered two more species, Cis curtidus Casey and Sulcacis lengi Dury, which also belong to this group. I have compared these with specimens determined as and fitting the descriptions of E. fronticornis, E. affinis, and E. bidentulus. C.curtulus is very similar to E. bidentulus, w^hile S. lengi is unique in having 9-segmented antennae, a somewhat shortened prosternum, and no sexual modifications on the head of the male. Since lengi is here selected as the type of Stdcacis Dury (1917), this genus becomes a junior synonym of Entypus. I would tentatively add R. japonicus and R. tokunagai, both from northern -Japan, to this genus, on the basis of descriptions and illustrations. I do not think the proposed division into subgenera is necessary, especially if it is based on the number of antennal segments.
There is one final matter that appears to have been overlooked by all of the workers in the Ciidae. This is the fact that the name Entypus was not available when used by Redtenbacher, having already been applied to a pompilid wasp (Hjanenoptera) by Dahlbom (1843). Entypus is a primary junior homonym and must be replaced. Of the two remaining available names which have been applied to the group, Sulcacis Dury is the oldest and is here considered as a replacement name. Although Sulcacis lengi Dury is the most atypical member of the group, it becomes the type species because of the priority of the generic name.
I do not think that S. lengi is distinct enough to be placed in a separate genus ; if this species were to be removed from the group, however, then Lohse ‘s Entypocis is the next oldest available generic name and E. bidentulus would be considered the type.

A synonymy and redefinition of Sulcacis is given below.
Sulcacis Dury, 1917, Jour. Cincinnati Soc. Nat. Hist., 22(2) :20; Leng,
1920:247. Type, by present designation, Sulcacis lengi Dury, 1917:21.
Eniypus Eedtenbaelier, 1847, Faun. Austr., (3):350 {non Dahlbom,1843:35); Thomson, 1863:193; Sahlberg, 1926:78; Lohse, 1964:118.
Type, by monotypy, Cis affinis Gyllenhal, 1827:628 (misidentified as Apate froniicornis Panzer).
Cis {Eniypus), Kiesenwetter, 1877:190; Eeitter, 1878:21.
Entypus (Eniypocis) Lohse, 1964, Ent. Blatter, 60(2) :121. Type, by original designation, Cis hideyitiilus Eosenhauer, 1847:58.
Cis (in part), Gyllenhal, 1827:624; Mellie, 1847:109; Mellie, 1848:236
Lacordaire, 1857:551; Jacquelin Du Val, 1861:237; Seidlitz, 1872:44
Abeille de Perrin, 1874:19; Eeitter, 1878:27; Seidlitz, 1891:282
Casey, 1898:78; Sehilsky, 1900 :37E; Eeitter, 1902:47; Dalla Torre,1911:5; Dury, 1917:5.
Ennearthron (in part), Mellie, 1848:360; Lacordaire, 1857:552; Jacquelin
Du Val, 1861:238; Abeille de Perrin, 1874:80; Eeitter, 1878:30;Sehilsky, 1900 :37B; Eeitter, 1902:59; Dalla Torre, 1911:23; Nobuchi,1960a: 41.
Ehopalodontus (in part), Jacquelin Du Val, 1861:238; Abeille de Perrin,
1874:76; Eeitter, 1878:30; Sehilsky, 1900 :37D; Eeitter, 1902:57;
Dalla Torre, 1911:21; Peyerimhoff, 1915:26; Nobuchi, 1960a: 39.
Form oblong to elongate, strongly convex and cylindrical ; vestiture short, usually dual, consisting of both erect and inclined bristles. Head moderately declined, partly covered by pronotum ;
frontoclypeal ridge of male usually with 2 small teeth or tubercles ; antennal fossa relatively deep. Antenna 9- or 10-segmented,with 3-segmented club; segments III and IV subequal, or III
longer than next 2 combined ; maxillary palp narrow and elongate. Pronotum somewhat constricted anteriorly, narrowly margined laterally; anterior edge usually simple in both sexes.
Elytra with relatively coarse, indistinctly dual, uniform or seriate punctation. Prosternum slightly tumid, concave laterally,slightly shorter than iutercoxal process which is narrow but not laminate. Protibia expanded at apex, outer apical angle forming a rounded process bearing several small spines. Metasternum convex, the suture short. Meso- and metatibiae slightly expanded and spinulose at apex. Sternite III of male with median pubescent fovea.
A small genus with species throughout the Holarctic region. Intermediate in its characters between Cis and Eridoulus, on the one hand, and Ceracis, Wacjaicis, and Malacocis on the other, differing from the former by the spinulose protibial apices and from LAWRENCE : CLASSIFICATION OF CIIDAE 279
the latter by the broader intercoxal process. The species which have been studied biolo<i’ically occur primarily on fungi of the Folyporns versicolor group (Paviour-Smith, 1960).
Two of Thomson’s genera, Iladraulc and Eridaulus, have been involved in considerable taxonomic confusion over the past 100 years. Lohse (1964), in his analysis of the problem, has referred
to it as a ” nomenklatorischen Tragikomodie,” an opinion with which I am inclined to agree.
Hadraide was originally proposed for the species Cis elongatulus Gyllenhal, characterized by the somewhat flattened body form, subquadrate prothorax, well-separated procoxae, and regularly
striato-punctate elytra. Seidlitz (1872), Kiesenwetter (1877), and Reitter (1878) considered it to be a subgenus of Cis. Schilsky (1900) first expanded the subgenus to include certain other species with a more or less parallel form, subquadrate prothorax, and elytra with rows of setae alternating with rows of large punctures. In addition to C. elongatulus (misspelled as elongatus in text, p. 59), he placed Cis setifer Reitter and Cis striatulus Mellie within Hadraule. Reitter (1902) continued the trend by including the following species as well : C. seriatopilosus Motschulsky, C. heiroglyphicus Reitter, C. hifasciatus Reitter, C. comptus Gyllenhal, C. coriaccus Baudi, and C. pumilio Baudi.
Jacobson (1915) doubtfully placed Cis fuscipcs Mellie, described from North America and Madeira, in the same group. Roubal (1936), having examined the type of C. dongatulus and discovered that it had 9-segmented, rather than 10-segmented, antennae, transferred the subgenus (including only the type
species) to the genus Enncarthron Mellie. Through a misunderstanding of the rules of nomenclature, Roubal replaced Hadraule with a new name, Knahlia. Lohse (1964) restored Hadraule and suggested that it be used in the original sense, as a separate genus including only the type species, Hadraule elorigafula.
Lohse further stated that Cis comptus and its relatives should be placed in the subgenus Eridaulus. This is entirely wrong, being based on an erroneous type designation, as will be explained below. The rest of the species mentioned above do seem to form a natural group, to which the American species Cis falli
Blatchley, Cis striolatus Casey, and Cis versicolor Casey might be added. H. elongatula shows a marked similarity to members of the C. comptus group and also to MapJioca hlaisdelli Casey,

latter differing mainly in the number of antennal club segments.
The genus Eridaulus Thomson (1863) originally included two species, Anohium nitidum Fabricius and Cis jacquemarti Mellie,which were characterized by the 10-segmented antennae, reflexed
“gular margin,” elytra with larger and smaller punctures (dual
punctation), short, carinate prosternum, and produced and dentate protibial apices. Seidlitz (1872), Sahlberg (1926), and a few others retained it as a genus, but most later authors reduced
it to a subgenus of Cis. Seidlitz added Cis glabra f us Mellie and C. lineatocrihratus Mellie, and Schilsky (1900) included C.quadridens Mellie as well. Roubal added one more European species, Cis hitu’berculosns Roubal, 1937 {=Cis hituherculatus Roubal, 1912, non Gorham), and in recent years Chujo (1940), Miyatake (1954), Nakane and Nobuchi (1955), and Nobuchi (1955, 1960b) have described several others from Japan. The only worker to designate a type for Eridauhis was Abeille de
Perrin (1874), who selected Cis comptus Gyllenhal. This species, however, was not originally included in the genus, and is therefore unavailable as a type. In the same work, Abeille de Perrin
designated C. nitidus as the type of Entypus, making a similar error. The name Eridaulus, then, is still available for the Cis nitidus group, contrary to the statement of Lohse (p. 119), and the type must still be designated. Since I have not examined the type specimens of either C. nitidus or C. jacquemarti, I hesitate to select one of them as the type at the present time.
In the past several years, I have examined specimens of Ciidae from various parts of the world, and it has become apparent to me that a number of other species described in Cis and the majority of those included in Xestocis Casey should be placed in Eridaulus. Casey (1898) proposed the genus Xestocis for 5 North American species with a carinate prosternum. Dury (1917), Brethes (1922), and Hatch (1962) added 14 more species, and most of those examined fall within this group.
Since a type has not been selected for Xestocis, I here designate X. levettei Casey, the best known of the American species. Miyatake (1954) noted the similarity of X. levettei to Cis (Eridaulus)
konoi Chujo, and I have noticed the resemblance of the former to C. nitidus, with respect to dual elytral punctation, carinate prosternum, dentate protibial apices, secondary sexual characters,
male genitalia, and larval urogomphi. I think there is little doubt that Xestocis should be considered a junior synonym of Eridaulus.


Since a number of species synonymies and new generic combinations are involved, I will not present here a complete list of American forms to be included in Eridaulus, but will
publish this separately in a forthcoming revision of the New World species. In addition to the American forms, several species of Cis from the Pacific region, such as Cis pacificus Sharp (Hawaii) and C. agariconae Zimmerman (Micronesia) should be placed in the genus.
Although Eridaulus appears to be quite close to the genus Cis, I think it should be given generic rank for several reasons.
According to Mayr, Linsley, and Usinger (1953), the size of the gap between two genera should be inversely proportional to the sizes of the groups involved. With its present constitution, the
genus Eridaulus is large enough, both in numbers of species and in extent of distribution, to warrant generic distinction. There are about 30 named species, and several more apparently undescribed, extending throughout the Holarctic region and as far south as New Zealand in the Pacific area. Furthermore, within the genus, several distinct subgroups can be recognized, and certain independent character trends can be followed. Finally, the group displays a biological unity, most of the species being associated with the larger, woody fruiting bodies of fungi such as Fomes and Ganoderma. This habit, which has been independently evolved in several unrelated genera, such as Ceracis and Xylographus, may be associated with structural modifications, such as the prosternal carination and convex form of the adult and the increased molar area in the larval mandible.
Paviour-Smith (1960) noted that the “headquarters” of E. nitidus was the fungus Ganoderma applayiatum, although the beetle was recorded from several other fungi, and this is quite similar
to the host range and headquarters of the American E. levettci.
Saalas (1923) reported E. quadridens, E. Uneatocribratus, and E. jacquemarti from Fomes pinicola, and the Japanese species E. rufocastaneus and E. nikkoensis have been taken on the same fungus. E. hiarmatus (Mannerheim) and several Eridaulus irom.
western North America occur on Fomes piuicola, F. annosus and related fungi. In Hawaii, where most of the Ciidae are collected by beating dead vines and branches, E. pacificus is the only species which has been reported feeding on the woody bracket fungi. It appears to me, then, that the members of this group represent a morphologically and biologically distinct genus, to which the name Eridaulus should be applied. Synonymies of Hadraule and Eridaulus and a redefinition of the latter genus are eiven below.


Hadraule Thomson, 1863, Skand. Col., 5:182; Sahlberg, 1926:71; Lohse,1964:119. Type, by monotypy, Cis elongaiulus Gyllenhal, 1827:627.
Cis {Hadraule), Seidlitz, 1872:44; Kiesenwetter, 1877:188; Reitter,1878:24; Seidlitz, 1891:281; Sehilsky, ]900:37F (in part); Reitter,1902:47 (in part); Dalla Torre, 1911:5; Jacobson, 1915:953 (in part).
Tityocis Peyerimhoff, 1918, Bull. Soc. Ent. France, 1918:141. Type, by monotypy, Pityocis coarctatus Peyerimhoff, 1918:142 ( = Cis elongatulus Gyllenhal). [See Peyerimhoff, 1933.]
Ennearthron {Knahlia) Roubal, 1936, Festsehr. 60 Geburtst. Embrik Strand,1:53. Type, by monotypy, Cis elongatulus Gyllenhal, 1827:627.
Hadraida Leng, 1920:246, incorrect subsequent spelling.
Eridaulus Thomson
Eridaulus Thomson, 1863, Skand. Col., 5:191; Seidlitz, 1872:45; Sahlberg,
1926:79. Originally included species: Anohium nitidum Fabricius,1792:238 and Cis jacquemarti Mellie, 1848:328. Type not designated.
Cis {Eridaulus), Kiesenwetter, 1877:191; Reitter, 1878:21, 24; Seidlitz,1891:281; Sehilsky, 1900 :37F; Reitter, 1902:48; Dalla Torre, 1911:5;Roubal, 1912:29; Roubal, 1937:39; Chujo, 1940:132; Miyatake,
1954:49; Nakane and Nobuchi, 1955:49; Nobuchi, 1955:56; Nobuchi,1960b :65.
Xestocis Casey, 1898, Jour. New York Ent. Soc, 6(2) :85; Dalla Torre,1911:20; Dury, 1917:15 (in part); Leng, 1920:247; Hatch, 1962:231.
Type, by present designation, Xestocis Icvettei Casey, 1898:85. NEWSYNONYMY.
Anohium, Ptinus, Cis auctt.
Form oblong, strongly convex ; vestiture of short, fine hairs, longer recumbent hairs, or short, stout setae. Head moderately declined, partly covered by pronotum ; f rontoclj’peal ridge of male produced on each side forming 2 flattened plates which are rounded to triangular ; genal ridge strongly elevated and carinate, forming relatively deep antennal fossa. Antennae 10-segmented, with a 3-segmented club. Pronotum strongly convex, variously margined laterally, anterior angles rounded to produced and acute ; anterior edge in male simple or produced and cmarginate. Elytra with dual punctation, consisting of larger nude macropunctures, and smaller setiferous micropunctures, the punctures uniform or seriate. Presternum relatively short, strongly tumid, concave laterally and carinate mesially ; intercoxal process relatively narrow but not laminate, subacute at apex. 

Protibia expanded at apex, outer apical angle produced and dentate. Sternite III of male with median pubescent fovea or patch.
A large genus with a primarily Holarctic distribution, but extending south at least in the Pacific region. Closely related to Cis and Strigocis, but distinguished by the strongly carinate prosternum, dual elytral punctation, and produced and dentate protibial apices. The genus differs from Hadraule, Orthocis, and Dulichocis in general body form and by the dentate protibial apices, as well as by the carinate prosternum. Most of the species in this genus feed on the larger woody fungi, such as Fomes and Ganoderma.
Casey (1898) proposed the genus Orthocis for the two American species, Cis puncfatus Mellie and Orthocis aterrima Casey,
which were distinguished from the species of Cis by the “…
more parallel form of the body . . . glabrous surface, margined elytral suture, and . . . simple apex of the anterior tibiae.”
Dury (1917) described a third species, 0. longida, from the eastern U.S. and Kraus (1908) added two species, 0. huesanus and 0. pidcher, from Florida. Another species, 0. platcnsis Brethes
(1922), was described from Argentina.

In the European literature, it has long been recognized that Cis alni Gyllenhal, and several related forms, such as C. perrisi Abeille de Perrin and C. coluhcr Abeille de Perrin, form a distinct group, characterized by the elongate body form, short pubescence, simple protibial apices, and lack of distinct sexual
modifications on the head or prothorax of the male. Lohse (1964) proposed for these species the subgenus Mcllieicis andselected Cis alni as its type. Although the exact limits of the subgenus were not given in Lohse ‘s paper, it was indicated that all species of Cis with rounded or truncate protibial apices should be included. Having studied the European literature and examined specimens in the American collections, I have come to the conclusion that both Cis punctatns Mellie sensu Casey and Orthocis aterrima Casey are very closely related to the Palearctic species in the Cis alni group. The name Orthocis, then, should apply to this group of species, and McUicicis should be considered a junior synonym.
In 1908, Kraus described 3 species of Ennearthron, E. annidatum, E. transversatum, and E. pallidum, from the southeastern United States and Cuba, which differ from the species of Orthocis only in the possession of 9-segmented antennae. I think it is only sensible to expand the definition of Orthocis to include these species as well, instead of leaving them in a genus which appears to be a conglomerate of unrelated forms. If the limits of the group are thus expanded, it becomes necessary to consider several other species which might be included. In 1919, Dury described the genus Dolichocis with the single species D. manitoha Dury, the main characteristics being the 9-segmented antennae, narrow, cylindrical form, vestiture of short bristles, and rounded protibial apices. The species differs from the American species of Orthocis, not only in its antennal segmentation, but by its stouter pubescence, anteriorly constricted prothorax, and the presence of 2 clypeal tubercles and a pubescent fovea on the head of the male. Hatch (1962) described a second species, D. incUstinctus, from western North America, which is very similar to and obviously congeneric with E7inearthron laricinmn (Mellie) of the European fauna. Other Palearctic species which might be in the same group are Ennearthron yuasai Chujo, E. pruinosiilum (Perris), and E. poriae Nakane and Nobuchi, the last two of which resemble Orthocis more than Dolichocis in the more quadrate prothorax with broader lateral margins and in the absence of sexual modifications on the head of the male. The problem is made more complex by the presence of 3 Palearctic species, Cis festivus (Panzer), Cis pygmaeus (Marsham), and Cis rhododactylus (Marsham), and one North American species, Cis angustus Hatch, which share certain characteristics with both Cis and Orthocis. Nyholni (1953) noted that the first three all have truncate or rounded protibial apices, and I have observed the same character in C. angustus.
Yet all of these have sexual ornaments on the head of the male, and they differ in general appearance from typical Orthocis. The setting of generic limits, if this is at all justified, presents a difficult problem, which can be solved only after a more detailed investigation of all species involved. In the present discussion, I will make a few suggestions based on a rather superficial study of a large number of forms from various parts of the world and a more intensive study of American representatives.
In the North American fauna, both Orthocis and Dolichocis seem to be well defined both morphologically and biologically.
All of the species of Orthocis are elongate and parallel, with a vestiture of very short and fine hairs, a fairly smooth and shining surface, margined elytral suture, flat or slightly tumid prosternum, rounded protibial apices, and an absence of sexual ornaments on the head of the male. There is a tendency in some species (0. longulus Dury) towards an extreme attenuation of
the body, and several subtropical forms are bicolored and may have 9-segmented antennae. Field observations indicate that at least some species occur under bark, in decaying branches and
vines, and apparently not in the tougher fruiting bodies of most Polyporaceae. The genus is more common in the southern part of the continent, and a number of unnamed species have been
seen from Central and South America. The two species of Bolichocis, on the other hand, dift’er in having the prothorax constricted anteriorly, the vestiture consisting of short, stouter bristles, and the head modified in the male. Both species occur in the northern part of the continent, and, like most Eridaulus, they occur mainly on the fruiting bodies of woody fungi.
In the European fauna, generic distinctions are not so easily made. Cis coluber Abeille de Perrin, C. reflexicollis Abeille de Perrin, and C. juglandis Reitter appear to be closely related to 0. alni, but they all have a stouter pubescence. Members of the Cis festivus group differ both in vestiture and in the presence of clypeal tubercles in the male. Of the species with 9-segmented antennae, E. laricinum undoubtedly belongs to Dolichocis, but E. priiinosulum appears to be closer to Orthocis. One character which has been overlooked by most European workers is the margined elytral suture in species of Orthocis. At the apex of each elytron, there is a distinct raised margin which curves laterad, leaving a narrow flat area between it and the edge of the suture. I have observed this in 0. alni, as well as in the American species. This character, in combination with others mentioned above, might serve to distinguish consistently the species of Orthocis from those of Cis and Dolichocis. Several other features which I have observed only in the North American Orthocis are the elongate prementum, elongate maxillary stipes with a flattened laterally placed laeinia, and the deeply emarginate, mesially unpigmented, 8th sternite of the male. An investigation of these and other more cryptic characters in the European species may shed some light on their true relationships.
Orthocis, like Eridaulus, is so similar to Cis that the question arises whether a generic distinction is justified or not. I think that the same criteria can be used here as were applied in the case of Eridaulus. The size of the group is quite large, especially in view of the number of unnamed species which I have seen,
and the distribution is probably cosmopolitan, with the majority of species occurring in tropical and subtropical regions. Several unique trends occur within the genus, and it appears that the group may have given rise to some of the peculiar endemic forms on the Hawaiian Islands and in other Pacific areas. The latter resemble GrfJiocis in the minute vestiture, flattened prosternum, simple protibial apices, and lack of male ornaments, but they vary considerably in body form, sculpture, and coloration. The trend towards extreme attenuation has already been mentioned.
The biological characteristics of the genus, although briefly mentioned above, deserve further comment. Most of the species of Ciidae feed on the sporophores of wood-rotting fungi, especially the more fibrous or woody fruiting bodies of the Polyporaeeae and Hydnaceae. These species have evolved various
morphological and physiological adaptations to enable them to cope with a substrate which not only presents a physical barrier to feeding but usually contains little moisture and a high percentage of pure chitin, unavailable to insects not possessing a symbiotic intestinal flora. In addition, there has been a tendency for various species to specialize in certain groups of host fungi (Paviour-Smith, 1960). The species of Ortliocis, whose habits have been recorded, appear to be general fungus feeders, occurring under bark, in dead branches and vines, in more humid situations, and in association with mycelia and fruiting bodies of a wide variety of wood-rotting fungi. The fungus sporophores are usually of a softer and more ephemeral type and their locations permit a certain amount of bacterial and fungal decomposition, so that the resulting substrate is more easily utilized by the beetles. Beniek (1952) has reported Cis alni from such diverse fungi as Exidia glandulosa (Tremellaceae), Stereum
rugosuni (Thelephoraceae), and Auricularia auricula-judae (Auriculariaeeae), while Lucas (1849) found the same species (as C. puncUdatus Lucas, non Gyllenhal) on Schizophyllum commune (Agaricaecae). Blatchley (1910, 1923) noted that 0. punctatus was taken by sifting debris from an oak log and
that 0. pulchcr was found by beating dead branches of oak.
Kraus (1908) bred Ennearthron fransvcrsatum from decaying rattan vines. Perris (1877) described the larva of Cis coluber from dead branches of chestnut and oak in which the fungus Thclcphora was oTowing; he also noted that the beetle could not be found on branches still remaining on the tree. In the same paper, he noted that the habits of C. alni and C. reflexicollis Abeille de Perrin were similar, and that C. ohlongus Mellie and C. pruinosulus were taken on fungus-infested elm branches. Zimmerman (1938, 1942) and Swezey (1954) have recorded a number of Pacific island species from dead vines and branches, and those which I have examined seem to fall within this same group.
One fact that makes the study of the particular group more difficult is that most of the shared characters are both simple and apparently primitive. Most of the special modifications which are characteristic of the family as a whole, such as the development of protibial structures for boring (expanded apices,
teeth, combs, spines), various prosternal modifications (shortening, carination, reduction of intercoxal process and increased coxal size), presence of sexual ornaments on the head and pronotum of the male, and the development of more complex punctation and vestiture, are all absent in the species of OrtJiocis. In addition, the generalized fungus-feeding habit may be considered primitive. Members of closely related families, such as the Lathridiidae and Corylophidae, share this habit with the species
of Orthocis, and Crowson (1955) thinks that this may have been characteristic of the ancestral Cucujoidea. If these characters are primitive or plesiomorphic, they are, according to Hennig
(1965), less reliable than derivative or apomorphic ones in determining relationships. If, on the other hand, Orthocis represents a collection of two or more convergent groups, which have
secondarily developed this type of feeding habit and thus have lost, through disuse, the structural modifications associated with boring, this convergence would probably be difficult to detect
because of the simplicity of the resulting characters. I favor the hypothesis that these characters are primitive, rather than derivative, and that the feeding habits probably represent those
of the common ancestor of the group. In certain other species, such as Hadraiile elongaiula and Maphoca hlaisdelli, which have similar biologies, this simplified condition also occurs, but other characters indicate that both of these are derivative forms, which may have evolved from a species in the Cis comptus group.
I think that most of the species discussed above should be placed in a distinct genus, to which the name Orilwcis applies.
The exact limits of the genus remain uncertain, especially with reference to the inclusion of Ch festivns and its relatives, and it is hoped that Dr. Lohse will shed some light on the relationships of these European species. I would definitely exclude Cis angustus Hatch, which does have a truncate protibial apex, but which occurs on Fomes pinicola, has clypeal tubercles in the male, and lacks a margin on the elytral suture.
The species of Dolichocis, although they share certain characters with Orthocis, should remain in a separate genus, differing by the absence of the margin on the elytral suture, the anteriorly
constricted prothorax, cylindrical form, and vestiture of short, stout setae. The genus should include the species laricinus (Mellie), yuasai (Chujo), mayiitoha Dury and indistinctus Hatch.
All four species occur on the larger, woody fruiting bodies of fungi, such as Fomcs pinicola and F. officinalis. Synonymies and redefinitions of Orthocis and DolicJiocis are given below.
Orthocis Casey
Orthocis Casey, 1898, Jour. New York Eiit. Soc, 6(2) :84; Kraus, 1908:77;
Dalla Torre, 1911:20; Dury, 1917:13; Leng, 1920:247; Brethes,
1922:302. Type, by present designation, Orthocis aterrima Casey,
Cis (Mellieicis) Lohse, 1964, Ent. Blatter, 60(2) :122. Type, by original
designation, Cis alni Gyllenhal, 1813:386. NEW SYNONYMY.
Enncarthron, Kraus, 1908:78.
Cis (in part), auctt.
Form elongate and somewhat depressed to narrowly elongate and cylindrical; vestiture of very short fine hairs or stouter bristles. Head slightly to moderately declined, only slightly covered by pronotum; frontoclypeal ridge without distinct sexual modifications in male ; antennal fossa shallow. Antenna 9- or 10-segmented, with 3- segmented club, segments III and IV usually elongate ; maxillary stipes elongate, laeinia lateral, palp relatively stout ; prementum elongate. Pronotum subquadrate, sides
narrowly margined to broadly margined and explanate, anterior angles truncate to slightly produced and rounded ; anterior edge simple in both sexes. Elytra usually parallel sided; punctation
single and relatively uniformly distributed; suture margined posteriorly, the margin curved laterad just before apex. Prosternum flat to slightly tumid, longer than intercoxal process which is fairly broad ; procoxae subtransverse, narrowly open behind. Protibia only slightly expanded at apex, outer apical
angle truncate or rounded. Metasternum slightly convex, the suture moderately long. Sternite III of male with a median pubescent fovea or patch.

A large, widespread g:emis, but mainly tropicopolitan. Closely related to Cis and Dolichocis, from which it differs by the rounded protibial apices, margined elytral suture, and lack of distinct sexual ornaments in the male. The species appear to be general fungus feeders, occurring often on dead vines and
branches penetrated by fungus mycelia.
Dolichocis Dury
Dolichocis Dury, 1919, Canad. Ent., 51:158; Hatch, 1962:234. Type, by
monotypy, Dolichocis manitoha Dury, 1919:158.
Cis (in part), Mellie, 1848:236; Lacordaire, 1857:551; Jaequelin Du Val,
1861:237; Seidlitz, 1872:44; Kiesenwetter, 1877:178.
Ennearthron (in part), Abeille de Perrin, 1874:80; Eeitter, 1878:30; Seidlitz, 1891:285; Schilsky, 1900 :37B; Eeitter, 1902:59; Dalla Torre,
1911:23; Chujo, 1941:85.
Form elongate, cylindrical; vestiture of short, stout, suberect bristles or squamae. Head moderately declined, partly covered by pronotum ; frontoclypeal ridge of male bituberculate, vertex
simple or foveate ; antennal fossa shallow. Antenna 9-segmented, with 3-segmented club ; maxillary stipes subquadrate, lacinia subterminal, palp relatively stout. Pronotum almost as long as wide, somewhat constricted anteriorly, sides narrowly margined, anterior angles not or barely produced ; anterior edge simple in both sexes. Elytra elongate and subparallel ; punctation single and fairly uniformly distributed. Prosternum slightly tumid, slightly longer than intercoxal process which is fairly broad and blunt at apex ; procoxae subtransverse, narrowly open behind.
Protibia only slightly expanded at apex, outer apical angle truncate or rounded. Metasternum slightly convex, suture moderately long. Sternite III of male with a median pubescent
A small genus restricted to the Holarctic region. Closely related to Cis and Orthocis, from which it differs in the rounded protibial apices, stout vestiture, constricted prothorax, and lack
of a margin along the elytral suture. The 4 species all occur on Fomes pinicola and related fungi.
Abeille de Perrin, Elzear
1874. Essai monographique sur les Cisides europeens et circamediter-
raneens. Camoin, Marseille. 100 pp.
Benick, L.
1952. Pilzkafer imd Kaferpilze. Acta Zool. Fennica, 70: l-l!50.
Blatchley, William S.
1910. All illustrated descriptive catalogue of the Coleoptera or beetles
(exclusive of the Ehynchophora) known to occur in Indiana.
Bull. Indiana Dept. Geol. and Nat. Ees., 1: 1-1385.
1923. Notes on the Coleoptera of southern Florida with descriptions
of new species. Canad. Ent., 55: 13-20, 30-36.
Brethes, Juan
1922. Descripcion de varios coleopteros de Buenos Aires. Ann. Soc.
Cient. Argent., 94: 263-305.
Casey, Thomas L.
1898. Studies in the Ptinidae, Cioidae, and Sphindidae of America.
Jour. NeAV York Ent. Soc, 6(2): 61-93.
Chujo, Michio
1940. Descriptions of a new and a rare species of the Japanese ciid-
beetles. Ins. Matsumurana, 14(4): 132-133.
1941. Description of a new ciid-beetle from Japan. Kontyu, 15(2) :
Ckowson, Eoy a.
1955. The natural classification of the families of Coleoptera. Lloyd,
London, 187 pp.
Dahlbom, Anders G.
1843. Hymenoptera Europae praecipue borealis . . . Tomus [1] : Sphex
in sensu Linnaeano. Luiidlierg, Lund, xliv + 528 pp.
Dalla Torre, K. W. von
1911. Cioidae. Pars 30. In W. Junk & S. Schenkling, edit., Coleopterorum Catalogus. Junk, Berlin, 32 pp.
Demarest, E.
1860. Tome 17. Coleopteres. Partie 3. In Jean C. Chenu, Encyclopedie d’histoire naturelle ou traite complet de cette science.
Didot, Paris, 360 pp.
Dury, Charles
1917. Synopsis of the coleopterous family Cisidae (Cioidae) of America north of Mexico. Jour. Cincinnati Soc. Nat. Hist.,
22(2): 1-27.
1919. A new ciside genus with new species from Manitoba. Canad.
Ent., 51 : 158.
Fabricius, Johann
1792. Entomologia systematica emendata et aucta. Tomus I [Pars 1].
Prost, Copenhagen, xx + 330 pp.
Gaubil, J.
1849. Catalogue synonymique des Coleopteres d ‘Europe et d ‘Algerie.
Maison, Paris, 296 pp.
Gyllenhal, Leonardo
1813. Insecta Svecica. Classis I. Coleoptera sive Eleuterata. Tomus I,
Pars III. Leverentz, Scaris, 730 pp.
1827. Tiiseeta Sveeiea, Classis I. Coleoptera sive Eleuterata. Tomus I.
Pars IV. Fleischer, Leipzig, 761 pp.
Hatch, Melville H.
1962. The beetles of the Pacific Northwest. Part III: Pselaphidae
and Diversicornia I. Univ. Washington Press, Seattle, ix +
503 pp.
Hennig, Willi
1965. Phylogenetic systematics. Ann. Eev. Ent., 10: 97-116.
Jacobson, G. G.
1915. Zhuki Eossii i zapadnoi Evropi. Euskovodstvo k opredeleniyu
zhukovi. Vip. XI. Devrient, Leningrad, pp. 865-1024.
Jacqublin Du Val, Pierre N. C.
1861. Manuel entomologique. Genera des Coleopteres d ‘Europe. Tome
3 [part]. DeyroUe, Paris, pp. 236-240, cat. p. 168.
1877. Anobiidae, Cioidae, Aspidophoridae. In W. F. Erichson, Naturgeschichte der Insecten Deutschlands. Abtheilung 1. Coleoptera. Band 5, Hiilfte 1. Moeser, Berlin, pp. 169-200.
Kraus, E. J.
1908. New bieolored Cioidae (Coleoptera). Proe. Ent. Soc. Washington, 10: 74-81.
Lacordaire, Jean T.
1857. Histoire naturelle des Inseetes. Genera des Coleopteres. Tome
4. Libr. Encycl. Eoret, Paris, 579 pp.

Leng, Charles W.
1920. Catalogue of the Coleoptera of America, north of Mexico.
Sherman, Mt. Vernon, N. Y., x + 470 pp.
Leng, Charles W. and A. J. Mutchlek
1927. Supplement 1919 to 1924 (inclusive) to Catalogue of the Coleoptera of America, north of Mexico. Sherman, Mt. Vernon, N. Y., 78 pp.
Lohse, Gustav A.
1964. Die in Mitteleuropa vertretenen Gattungen der Cisidae (1.
Beitrag zur Kenntniss der mitteleuropaischen Cisidae). Ent.
Blatter, 60(2): 116-122.
Lucas, H.
1849. Exploration scientifique de I’Algerie, Zoologie. Histoire naturelle des animaux articules. 2 Partie. Inseetes. Imprim. Nat., Paris,
590 pp.
Mayr, Ernst, E. G. Linsley and E. L. Usinger
1953. Methods and principles of systematic zoology. McGraw-Hill,
New York, vii + 328 pp.
Mellie, J.
1847. Melanges et nouvelles. Eev. Zool. Cuvierienne, 10: 108-110.
1848. Monographie de 1 ‘ancien genre Cis des auteurs. Ann. Soc. Ent.France, Ser. 2, 6: 205-274, 313-396, pis. 9-12.
1954. studies on the Japanese Ciidae, I. (Coleoptera). Sei. Eep.
Matsuyama Agr. Coll., 14: 40-67, 11 pis.
Nakane, Takehiko and Akira Nobuchi
1955. On a new genus and six new species of ciid-beetles from Japan(Ciidae, Coleoptera). Sci. Rep. Saikyo Univ., Ser. A, 2(1):
Nobuchi, Akira
1955. Studies on the ciid-beetles from Japan. (I) With descriptions
of a new genus and some new species (Ciidae, Coleoptera). Ent.Rev. Japan, 6(7) : 53-58, pi. 12.
1960a. Ciidae from Hokkaido (Coleoptera). Ent. Rev. Japan, 11(2):
37-41, 1 pi.
1960b. On a new species of Cis Latreille from Japan (Coleoptera,Ciidae). Akitu, 9: 65-66.
Nyholm, Tord
1953. Bemerkungen iiber einige schwedische Cis-Artcn (Col., Ciidae).
Ent. Tidskr., 74: 187-196.
Paviour-Smith, Kitty
1960. The fruiting-bodies of macrofungi as habitats for beetles of the
family Ciidae (Coleoptera). Oikos, 11(1): 43-71, 1 pi.
Perris, Edouard
1877. Larves de Coleopteres [part]. Ann. Soc. Linn. Lyon, N.S., 23:
1-43, 14 pis.
Peyerimhoff, p. de
1915. Notes sur la biologie de quelques Coleopteres phytophages du
Nord Africain (Deuxieme serie). Ann. Soc. Ent. France, 84:
1918. Nouveaux Coleopteres du Nord-Africain (Vingt-huitieme note:
Faune du Pin d’Alep). Bull. Soc. Ent. France, 1918: 141-143.
1933. Les Coleopteres attaches aux coniferes dans le nord de 1 ‘Af rique.
Ann. Soc. Ent. France, 102: 359-408.
Redtenbacher, Ludwig
1847. Fauna Austriaca. Die Kafer [Part 3]. C. Ceroid, Vienna, pp.
Reitter, Edmund
1878. Nachtrage und Erganzungen zur Bearbeitung der Cioiden von H. V. Kiesenwetter. Deutsch. Ent. Zeitschr., 22(1): 21-30.
1902. Analytische Uebersicht dcr palearctischen Gattungen und Artcn der Coleopteren-Familien: Byrrhidae (Anobiidae) und Cioidae.
Verhandl. Naturforsch. Ver. Briinn, 40: 1-61.
Rosenhauer, Wilhelm G.
1847. Beitriige zur Insekten-Fauna Europas. Erstes Bandchen. Blaesing, Erlangen, x + 159 pp.
RouBAL, Jan
1912. Koleopterologicke vysledky me cesty na Kavkaz v cervenci r.1910. Casopis, 9: 25-28,124-26.
1936. Besehreibung einer neuen Untergattung, einer Varietat und einer Form der Coleopteren. Festschr. 60 Geburtst. Embrik
Strand, 1 : 53-58.
1937. Koleopterologische Notizen. Ent. Nachrichtenblatt, 11: 38-40.
Saalas, Uunio
1923. Die Fichteiikafer Filmlands. II. Spezieller Teil 2 und Larven-bestimmungstab. Suom. Tiedeakat. Toimituskia, Sarja A, 22
(1): 1-746, 28 pis.
Sahlberg, John
1926. Enumeratio Coleopteroruni Fenniae. Lamellicornes, Platysoma,
Xylophagi, Fungicola. Ann. Soe. Zool.-Bot. Fennicae, 4(1):
1900. Heft 37 in H. C. Kuster, G. Kraatz, and J. Schilsky, Die Kiifer Europas. Bauer & Raspe, Niirnberg, pp. i-iv, 1-100, A-W.
Seidlitz, Georg
1872. Fauna Baltica. Die Kafer (Coleoptera) der Ostsee-Provinzen Russlands. Laakmann, Dorpat, xlii + 142 pp.
1891. Fauna Transylvanica. Die Kafer (Coleoptera) Siebenbiirgens.
Hartung, Kaliningrad, Ivi + 192 + 914 pp.
SwEZEY, Otto
1954. Forest entomology in Hawaii. Spec. Publ. Bernice P. Bishop Mus., No. 44, ix + 266 pp.
Thomson, C. G.
1863. Skandinaviens Coleoptera, synoptiskt bearbetade. Tome 5.
Lundberg, Lund, 340 pp.
Wankowicz, Jean
1869. Notices entomologiques. Ann. Soe. Ent. France, Ser. 4, 9: 411-422.
Zimmerman, Elwood C.
1938. Ciidae of southeastern Polynesia (Coleoptera). Occ. Pap.
Bernice P. Bishop Mus., 14: 199-217.
1942. Ciidae of Guam. In Insects of Guam — I. Bull. Bernice P.
Bishop Mus., 172: 47-52.
(Received 5 March 1965.)

Leave a Reply

Your email address will not be published. Required fields are marked *