The Ciidae (Coleoptera) of New Brunswick, Canada: New records and new synonyms

Abstract The Ciidae of New Brunswick, Canada are reviewed. Seventeen species are recorded for New Brunswick, including the following 10 species that are newly recorded for the province: Ceracis singularis (Dury), Ceracis thoracicornis (Ziegler), Cis angustus Hatch, Cis fuscipes Mellié, Cis horridulus Casey, Cis striatulus Mellié, Dolichocis laricinus (Mellié), Malacocis brevicollis (Casey), Orthocis punctatus (Mellié), and Plesiocis cribrum Casey. Additional locality data are provided for the following species previously known from the province: Cis americanus Mannerheim, Cis creberrimus Mellié, Cis levettei (Casey), Cis submicans Abeille de Perrin, Dolichocis manitoba Dury, Hadreule elongatula (Gyllenhal), and Octotemnus glabriculus (Gyllenhal). Seven synonyms are proposed here; Cis pistoria Casey with Cis submicans Abeille de Perrin; Cis fraternus Casey, Cis macilentus Casey and Cis striolatus Casey with Cis striatulus Mellié; Dolichocis indistinctus Hatch with Dolichocis laricinus (Mellié); and Octotemnus denudatus Casey and Octotemnus laevis Casey with Octotemnus glabriculus (Gyllenhal). Lindgren funnel traps provided the majority of specimens for 15 of the 17 species reported from New Brunswick and were the sole source of specimens for seven of the 10 species newly reported here, suggesting they are a very useful tool for sampling Ciidae in the forests of New Brunswick.


Introduction
The systematics, taxonomy, and biology of the North American Ciidae (minute treefungus beetles) are well known as a result of the works by Lawrence (1965Lawrence ( , 1967Lawrence ( , 1971Lawrence ( , 1973Lawrence ( , 1982 and Thayer and Lawrence (2002). Adults and larvae live in and feed on basidiomes of various species of basidiomycete fungi, commonly known as polypores or bracket fungi, with most ciid species limited to a few host species (Lawrence 1973, Thayer and Lawrence 2002, Orledge and Reynolds 2005. The Ciidae of the Maritime Provinces of Canada were reviewed by Majka (2007). He provided an overview of the history of collecting, host usage of species occurring in the region, and a discussion on the distribution and zoogeography of the fauna. Fifteen species were reported from the region, including six species from New Brunswick. Cis pistoria Casey was newly recorded from New Brunswick. Majka (2007) noted that sampling in New Brunswick was inadequate and that additional species would likely be documented with more intensive sampling.
During a study in New Brunswick to develop tools for improved detection of invasive species of Cerambycidae, many Ciidae were collected from Lindgren funnel trap samples. Other specimens were collected during general sampling, and additional material was found in several museum collections. Ten species new to New Brunswick were found during this survey as well as many additional records of species previously known from the province. The purpose of this paper is to document these records and to present seven new synonyms.

Methods and conventions
Collection methods. The following records are based, in part, on specimens collected as part of a general survey to document the Coleopteran fauna of New Brunswick. Other records were obtained from specimens contained in the collections listed below. Most species records came from samples collected from Lindgren funnel traps deployed at 27 sites  traps per site) between 2009 and 2015. At many sites, starting during 2012, traps were deployed in the upper canopy as well as in the understory, usually in equal numbers, although at a few sites, only canopy traps or understory traps were used. Canopy traps were 10-20 m above the ground, whereas understory traps were 1-1.5 m above the ground (i.e., 30-50 cm from the bottom of the collecting cup to the ground). In both cases, traps were suspended from a rope such that the trap was at least 1 m from the main stem of trees and at least 30 m from another trap. Traps were baited with various combinations of lures for detecting Cerambycidae. However, data on attractants were not collected for the Ciidae. For details of the methods used to deploy Lindgren traps, for sample collection, and lure combinations used, see Webster et al. (2012aWebster et al. ( , 2012b, Hughes et al. (2014), and Webster et al. (2016). Locality and habitat data are presented as on labels for each record. Two labels were used on many specimens, one that included the locality, collection date, and collector, and one with macro-and microhabitat data and collection method. Information from the two labels is separated by a // in the data presented from each specimen.
Distribution. Every species is cited with current distribution in Canada and Alaska, using abbreviations for the state, provinces, and territories. New records for New Brunswick  Taxonomy. Specimens were initially separated into morphospecies and then identified using keys to North American Ciidae provided by Lawrence (1967Lawrence ( , 1971. They were also carefully compared with identified specimens deposited in the Coleção Entomológica do Laboratório de Sistemática e Biologia de Coleoptera, Universidade Federal de Viçosa, Viçosa, Minas Gerais, Brasil (CELC) and with material borrowed from North American and European museums, which included material compared with types, collected in type localities, and identified by former and current specialists on Nearctic and Palaearctic ciids (e.g., Adolf Lohse, Alexander Kompantsev, Glenda Orledge, Johannes Reibnitz, John Lawrence, Makoto Kawanabe, among others). As the North American Ciidae fauna were completely and carefully revised by Lawrence (1971), who examined most types, proposed synonyms, and redelimited species, identification of North American species is reliable, and the morphological limits of most species are well understood. The major problems that persisted in the taxonomy of North American ciids are also mentioned and discussed in that work, mainly regarding a few possible cases of conspecificity between North American and European species. In order to check these, the ciid species from NB were compared with Holarctic material and data in literature (e.g. Lohse 1967, Kawanabe 2002. When necessary and possible, males were dissected, and their genitalia mounted on slides following the protocol provided by Lopes-Andrade (2011). A complete list of references and original combinations is provided only for species with new synonyms proposed here.
It is important to emphasize that we propose new synonyms only for species with well-known morphological limits and that were previously studied by authors who conducted faunistic or revisionary works on the North American or the European Ciidae faunas. The morphological limits of these species are well established in literature, there is available material deposited in museums and used for comparison, and there remains no doubt on their identification. For instance, there is no doubt that North American specimens currently identified as Octotemnus laevis all refer to a single species, and its definition was revised previously by Lawrence (1971). There is no doubt that European and Asian ciids identified as O. glabriculus all refer to a single species, and the unique species that could be confounded with it, O. rugosopunctatus Drogvalenko, was also examined. But there is robust evidence that O. laevis and O. glabriculus refer to a single, Holarctic species, and thus it was necessary to resolve this so as not to perpetuate the problem. All other synonyms proposed here follow this same reasoning. In the case of Orthocis punctatus, a cryptic species for which we do not have enough material identified by renowned specialists or compared with type specimens, we have retained the name used by Lawrence (1971). However, it is likely that further synonyms will arise as relevant material is examined.
Some species names cited here were proposed by Mellié in a work published in two separate parts, and in the last decades, there has been much confusion in the literature regarding the publication year of his monograph. Authors have cited both parts of Mellié's monograph as being published in either 1848 or 1849, or 1848 for the first part and 1849 for the second. Here, we used 1849 as the publication date for both parts of Mellié's work, following Orledge and Booth (2006) who stated "(...) it is clear that pages 313-396 [part 2] were published in April/May 1849, and that pages 205-274 [part 1] may well have been published in January 1849." A more detailed discussion on this matter can be found in Orledge and Booth (2006), but it is important to emphasize here that subsequent authors are following their suggestion (e.g., Jelínek 2007, Reibnitz et al. 2013, Sandoval-Gómez et al. 2014.

Species accounts
Species with a † are adventive to Canada, species with a * are Holarctic. The determination that a species was a new record was based on information in the print version of Bousquet et al. (2013). Four of the species found in NB are new junior synonyms of four Palaearctic names, respectively: Cis pistoria Casey with C. submicans Abeille de Perrin; C. striolatus Casey with C. striatulus Mellié; Dolichocis indistinctus Hatch with D. laricinus (Mellié); and Octotemnus laevis Casey with O. glabriculus (Gyllenhal). However, in two cases, the junior synonyms proposed here already had previously proposed synonyms to them: C. fraternus Casey and C. macilentus Casey, previously synonymized to C. striolata; O. denudatus Casey, previously synonymized to O. laevis. These were well-established synonyms, and we also include these names as new synonyms of C. striatulus and O. glabriculus, respectively. We follow recent alterations on a few names of European species provided by Jelínek (2007).

Family Ciidae Leach, 1819 Subfamily Ciinae Leach, 1819 Tribe Ciini Leach, 1819
Ceracis singularis (Dury, 1917), new to New Brunswick Distribution in Canada and Alaska. ON, QC, NB . All new records of Ceracis singularis (Dury) in NB were based on specimens captured in Lindgren funnel traps or a flight intercept trap. This species is widespread in the province (seven localities) but was captured in low numbers at each site. This species was found in hardwood, mixed, and pine forests. These are the first records of this species from the Maritime Provinces of Canada.  . All new records of Ceracis thoracicornis (Ziegler) in NB were based on specimens captured in Lindgren funnel traps. This species is widespread in the southern half of the province but was found at only five localities and in relatively low numbers. This species was found in hardwood, mixed, and pine forests.  . Nearly all records of Cis americanus Mannerheim, which is widespread (12 sites) in NB, were based on specimens captured in Lindgren funnel traps from various forest types. The first records of this species from NB were specimens reared from Piptoporus betulinus (Bull.) Fr. (=Polyporus betulinus) (birch polypore) (Polyporaceae) from two localities (Wirral and Ludlow) (Pielou and Verna 1968 Distribution in Canada and Alaska. BC, NB . All new records of Cis angustus Hatch from NB were based on specimens captured in Lindgren funnel traps. This species is widespread (seven localities) in the province but was captured in low numbers at all sites. Adults were captured mostly in conifer-dominated forests and mixed forests. These are the first records of this species from eastern Canada. According to Lawrence (1971), C. angustus is restricted to coniferous forests at higher elevations in the mountains of the Pacific coast of southern BC, south to the southern Sierra Nevada in CA. He mentions further field work might reveal a broader distribution and that this species is part of the Holarctic faunal element. Our records in NB lead to an apparent disjunct distribution of the species in Canada, However, it is likely with additional field work that this species will be found in the intervening areas of northern Canada.

Cis americanus Mannerheim, 1852
Taxonomic notes. A male was dissected, and its determination as C. angustus was confirmed by John Lawrence. This species was also compared with the closely related Palaearctic species Cis fagi Waltl and Cis fusciclavis Nyholm, and it was confirmed that C. angustus is distinct from those two species.

Cis creberrimus Mellié, 1849
Available data. New Brunswick, Westmorland Co., Moncton, 13.VII.1987, P. Maltais (UMC) Distribution in Canada and Alaska. ON, QC, NB, NS ). Cis creberrimus Mellié was first reported from NB by Bousquet et al. (2013) but without supporting data. Chris Majka supplied the data for the specimen, which is in the collection of the University of Moncton. However, the curator of the collection was unable to locate the specimen. The inclusion of this species in the fauna of NB should be considered provisional until the specimen can be located.
Taxonomic note. Cis creberrimus is somewhat similar to C. angustus and may be confounded with this species. In view of this, the NB specimen identified as Cis creberrimus, could be C. angustus.  . Nearly all of the new records of Cis fucipes Mellié in NB were based on specimens captured in Lindgren funnel traps. This species is widespread (25 sites) and the most common ciid in the province, being captured in traps at nearly all sites and forest types where Lindgren traps were used. All new records of Cis horridulus Casey from NB were provided by specimens captured in Lindgren funnel traps. This species is widespread (10 localities) in the province. Adults were captured in hardwood, mixed, and conifer forests.

Cis horridulus Casey, 1898, new to New Brunswick
Taxonomic notes. A male was dissected, and its genitalia compared with those of the closely related Palaearctic species C. punctulatus Gyllenhal and C. tomentosus Mellié, and it was confirmed that C. horridulus is not conspecific with these two species.   . Most records of Cis levettei (Casey) in NB were from specimens captured in Lindgren funnel traps. This species is widespread (13 localities) and fairly common in the province, occurring in hardwood, mixed, and conifer forests. Specimens with habitat data were found in Piptoporus betulinus, Fomes fomertarius (L.) Fr., Polyporus varius (Pers.) Fr., and bracket fungi. This species was first reported from NB by McNamara (1991) but without supporting data (many specimens in CNC that are reported here).

Cis levettei (Casey, 1898)
Taxonomic notes. A male was dissected, and its genitalia compared with those of the closely related Palaearctic species C. castaneus (Herbst), Cis glabratus Mellié, C. hanseni Strand, C. jacquemartii Mellié, and C. lineatocribatus Mellié, and it was confirmed that C. levettei is not conspecific to any of these.  . All new records of Cis striatulus Mellié from NB were based on specimens captured in Lindgren funnel traps. This species is widespread (16 localities) and fairly common in the province, occurring in hardwood, mixed, and conifer forests.

Cis striatulus
Taxonomic notes. A drawing of the male tegmen of the North American C. striolatus Casey was provided by Lawrence (1971), who also commented that the species would be a synonym of C. striatulus, the latter occurring in Central Europe, the Caucasus, and northern Africa. The type locality of C. striolatus is Salida (Colorado). Although no specimen collected near the type locality was examined, the identification of the species is confident, and sufficient data on its morphological limits were provided by Lawrence (1971). The type locality of C. striatulus is southern France, and there were specimens from localities in France, Germany, and northern Iran (the Caucasus) available for comparison. A few males were dissected, confirming that tegmen shape is the same as shown in Lawrence (1971). External morphological features and known intraspecific variation matched between Nearctic and Palaeartic populations of C. striolatus and C. striatulus, respectively, so we propose their synomymization here. Cis striolatus had two junior synonyms previously proposed by Lawrence (1971): C. fraternus Casey and C. macilentus Casey. These names refer to variations of C. striolatus, and thus, we also propose them as new synonymies of C. striatulus. As Cis flavipes Lucas, 1847 is a junior homonym of Cis flavipes Motschulsky, 1845, the oldest available name becomes Cis striatulus Mellié, 1849.  . Most records of C. submicans from NB were based on specimens captured in Lindgren funnel traps. This species is widespread (nine localities) and fairly common in the province, with most records from hardwood-dominated and mixed forests. Some specimens were found under bark of a log covered with polypore fungi. Cis submicans (as Cis pistoria Casey) was first reported from NB by Majka (2007) on the basis of a specimen from the Meduxnekeag Valley Nature Preserve (record is reported here).

Cis submicans
Taxonomic notes. Cis pistoria is the only New World member of the Cis boleti species group, which also includes the Palaearctic C. boleti (Scopoli), C. micans (Fabricius), C. polypori Chûjô (also treated as a subspecies of C. boleti), C. rugulosus Mellié and C. submicans Abeille de Perrin (Chûjô 1939, Lawrence 1971. Specimens of all these species and subspecies were available for comparison and, among them, aedeagi of C. pistoria from North America and those of C. submicans were indiscernible. Cis pistoria was described by Casey (1898) based on specimens from "Rhode Island (Boston Neck)" in USA, a locality on the northeastern coast and about 500 linear km south of NB. Most reports of C. pistoria to date were in northeastern USA and in Canada (Lawrence 1971. Cis submicans was described by Abeille de Perrin (1874) as a variety of Cis setiger Mellié (currently C. villosulus) and based on specimens from the Caucasus and Poland. It is important to note that most C. submicans available for comparison were also from northern Iran (in the Caucasus) and Poland. As specimens currently recognized as Cis pistoria fits well in the morphological limits of the Palaearctic Cis submicans, we propose their synonymization here. Cis submicans has a Holarctic distribution, occurring in Europe and the Caucasus, and in northeastern North America. Fig. 10 Ennearthron laricinum Mellié, 1849: 355, pl. 12, fig. 3 Dolichocis indistinctus Hatch, 1962: 234, new synonym Material examined. New Brunswick, Kent Co., Kouchibouguac National Park, 46.8072°N, 64.9100°W, 12-14.VI.2015, 7-22.VII.2015 Alderson & V. Webster // Jackpine forest, Lindgren funnel traps, 1 m high (1, AFC; 1, RWC). Northumberland Co., ca, 2.5 km W of Sevogle, 47. 0876°N, 65.8613°W, 28.V-11.VI.2013, 11-26.VI.2013, 10-24.VI.2014 Distribution in Canada and Alaska. BC, QC, NB . All new records of Dolichocis laricinus (Mellié) from NB were based on specimens captured in Lindgren funnel traps. This species is currently known from three localities in NB. Specimens were captured in jack pine forests (Pinus banksiana Lamb.) at two localities and a mixed forest. These are the first records of this species from the Maritime Provinces.

Dolichocis laricinus (Mellié, 1849)*, new to New Brunswick
Taxonomic notes. The genus Dolichocis Dury has only four species: D. indistinctus Hatch and D. manitoba Dury from North America, the Eurasian D. laricinus (Mellié), and D. yuasai (Chûjô) from Japan. The possible synonomy of D. indistinctus Hatch and D. laricinus was first proposed by Lawrence (1971). The type locality of Dolichocis indistinctus (Stanley, BC) is close to the northwestern coast, but the species has a broad distribution in North America, and there is currently no doubt about its identification. The type locality of D. laricinus is Paris (France), and specimens from Poland and France were available for comparison. A male D. indistinctus from NB was dissected, and its aedeagus is exactly the same as in European specimens of D. laricinus. Specimens of D. manitoba were also examined and dissected, confirming that its male genitalia are quite distinct from those of D. indistinctus and D. laricinus. Unfortunately, no D. yuasai were available, and no opinion can be given on this species. As D. indistinctus is well within the morphological limits of D. laricinus, we propose their synonymy. The species has a Holarctic distribution. In North America, it occurs on the same host fungi of D. manitoba Dury, but D. laricinus appears to be much rarer (Lawrence 1971). Fig. 11 Material examined. New Brunswick, Albert Co., Fundy N.P., Point Wolfe R. Trail, 25.VII.1968, E.E. Lindquist, Ex: bracket fungi (8, CNC). Northumberland Co., Ludlow, 18.VII.1967, D. P. Pielou, Ex: Polyporus betulinus, H-21 (91, CNC); Taxis [River], 26.V.1967, D.P. Pielou, Ex: Polyporus betulinus, G-364 (750, CNC); ca. 2.5 km W of Sevogle, 47. 0876°N, 65.8613°W, 9-23 (Casey) 14 Orthocis punctatus (Mellié)   . Most records of D. manitoba from NB were based on specimens captured in Lindgren funnel traps. This species is widespread (12 localities) and fairly common in the province. This species was first reported from NB by Pielou and Verna (1968) (records included above). Dolichocis manitoba was captured in hardwood, mixed, and conifer forests. Specimens were reared from Piptoporus betulinus (=Polyporus betulinus) from four sites by D.P. Pielou. Fig. 12 Material examined. New Brunswick, Northumberland Co., Ludlow, 6.VI-31. VII.1967, 21.VI.1968, 2.VII.1968 19-25.V.2009, 2-9.VI.2009, 9-16.VI.2009, 16-24.VI.2009 Distribution in Canada and Alaska. NB . Hadreule elongatula (Gyllenhal) is widespread throughout Europe, Siberia, and North Africa (Lawrence 1971). Lawrence (1971) hypothesized that the species would have a broader distribution in North America, but in the subsequent decades, it has been found only in NB, where it was first reported by Pielou and Verna (1968) from specimens reared from Piptoporus betulinus (=Polyporus betulinus) at two localities (reported above). It was probably introduced from Europe (Lawrence 1971). All recent records of this species from NB were based on specimens captured in Lindgren funnel traps. This species is widespread (seven localities) and fairly common in the province. This adventive species was captured mostly in conifer forests in NB.

Hadreule elongatula (Gyllenhal, 1827) †
Taxonomic notes. It is worth mentioning that the correct spelling of the genus name is Hadreule, not Hadraule (see Orledge and Booth 2006). In several works, the publication date of this genus, described by Thomson, was cited as being 1863. In Thomson (1863) the spelling is "Hadraule". However, the name was indeed proposed four years before by the same author, but spelled as "Hadreule" (Thomson 1859: 91). Therefore, "Hadraule" is an incorrect subsequent spelling. Fig. 13 Material examined. New Brunswick, Carleton Co., Jackson Falls, "Bell Forest", 46. 2200°N, 67.7231°W, 12-19.VI.2008, 19-27.VI.2008 thocis punctatus is widespread (21 localities) and one of the most common species of Ciidae in NB. Specimens were captured in hardwood, mixed, and conifer forests.

Malacocis brevicollis (Casey, 1898), new to New Brunswick
Taxonomic notes. There is evidence that Orthocis punctatus comprises at least two species in North America and that they may be conspecific to European species. This problem was first noted by Lawrence (1971), but he kept all known forms under the same name due to the lack of a comparative study of male genitalia. The matter is beyond the scope of this project, and we prefer to attribute the name Orthocis punctatus to the species from NB. Distribution in Canada and Alaska. BC, AB, MB, QC, NB . All new records of Plesiocis cribrum Casey from NB were based on specimens captured in Lindgren funnel traps. This species is currently known from six localities in NB and appears to be uncommon. Specimens were captured in hardwood, mixed, and conifer forests. These are the first records of this species from the Maritime Provinces.