The identity of Parandra laevis Latreille , 1804 and nomenclatural changes in the Parandrinae ( Coleoptera , Cerambycidae )

Attelabus glaber (DeGeer, 1774) currently the type species of Hesperandra (Zikandra) Santos-Silva, 2003 is considered the type species of Parandra Latreille, 1802 (here validated). Hesperandra Arigony, 1977 and Hesperandra (Zikandra) become junior synonyms of Parandra Latreille, 1802, and Gnathophorus Kirby, 1837 is also confi rmed as a synonym of this genus. Birandra Santos-Silva, 2002 is used to replace Parandra sensu lato. Parandra laevis Latreille, 1804 is placed as a synonym of Attelabus glaber DeGeer, 1774. New names are chosen to replace Parandra laevis Schönherr, 1817 (= sensu auctorum) (redescribed here) and the subgenus Parandra sensu Santos-Silva, 2002, respectively: Birandra (Yvesandra) latreillei and Birandra (Yvesandra). Protospondylis Linsley, 1942 is revalidated, excluded from Parandrinae, and placed incertae sedis in the Cerambycidae. An updated list of the reallocated species is given.


Introduction
Th e purpose of this paper is to resolve confusion regarding the identity of Parandra laevis Latreille, 1804 and to propose a new generic name for the species currently attributed to Parandra Latreille, 1804 (Monné 2006).
Parandra laevis Latreille, 1804 currently placed in Parandra (Parandra) (Monné 2006), is regarded as the type species of the genus Parandra Latreille, 1802(Th omson 1864).Bousquet (2008) recognized the fact that P. laevis Latreille, 1804 was, under the current Code (ICZN 1999), a junior objective synonym of Attelabus glaber DeGeer, 1774.Attelabus glaber is the type species of Parandra Latreille, 1802, by monotypy and currently placed in Hesperandra (Zikandra) Santos-Silva, 2003(Monné 2006).Th us any genera/subgenera which have the type species A. glaber become synonyms of Parandra.Th is also includes the mainly forgotten genus Gnathophorus Kirby, 1837(Bousquet 2008).Bousquet (op. cit.) correctly points out that the species Parandra laevis sensu auctorum is without valid synonyms (Monné 2006;Santos-Silva 2002) and thus a new name is needed.He also proposed that if it was deemed desirable to retain the species concept of P. laevis Latreille, 1804, to maintain current taxonomic stability, that it would be necessary to apply to the commission to suppress Olivier (1803) as the author of Parandra in favour of Latreille (1804).Th e problem with this suggestion is that Latreille's use of the name "laevis" in 1804 does not defi ne or constitute a species separate from Attelabus glaber DeGeer, 1774 (see below) and as such cannot be used to validate the genus Parandra in Latreille (1804).It is therefore not an acceptable option.Th e subsequent validation, in this paper, of Latreille, 1802 as the author of the genus Parandra does not alter the need to propose a new generic name for the species currently placed in Parandra Latreille, 1804 (Monné 2006) as Attelabus glaber DeGeer (1774) is the only species given in the 1802 text.Th us Hesperandra Arigony, 1977 becomes a junior synonym of Parandra Latreille, 1802.Parandra laevis as a separate species concept, as proposed by Schönherr (1817) and Gyllenhal (1817), is retained and renamed accordingly.

Materials and methods
We examined specimens, males and females, of all species (including non American) in the same subgenus as Parandra (Parandra) laevis Latreille, 1804, specimens of Hesperandra (Zikandra) glabra (DeGeer, 1774) from various countries, and specimens of Neandra brunnea (Fabricius, 1798).Th ese specimens belong to several museums and private collections, and were used to establish the identity of Parandra laevis Latreille (1802Latreille ( , 1804Latreille ( , 1806Latreille ( , 1807Latreille ( , 1818) ) and Olivier (1803) as it is understood today.Th e generic name Parandra, formally attributed to Latreille (1804), was subsequently attributed to Olivier (1803) (Bousquet 2008).However the concept of the genus Parandra was fi rst proposed for the species Attelabus glaber DeGeer, 1774 by Latreille in 1802:160 (noted by Th omson, 1858, and Lameere, 1902).In the 1802 text Latreille used the vernacular name "Parandre; parandre".As the name was not given a Latin form, this 1802 description of Parandra has been considered to be invalid according to ICZN Code, 1999:109.However the failure to give a Latin form for the name would appear to have been an error as all other genera mentioned in the Latreille (1802) text have the second expression of the name in Latin.According to Article 32.5.1 (ICZN Code 1999:39) "if there is clear evidence of an inadvertent error, such as a lapsus calami or a copyist's or printer's errors, it must be corrected".In accordance with the format of the 1802 text the corrected spelling of the genus "Parandre; parandre" is "Parandre; Parandra", and thus, the generic name Parandra becomes available from Latreille's 1802 publication (Dr.Yves Bousquet is in agreement with this conclusion, pers.comm.).
It was common practice in the early 19 th century for European authors, particularly in the dictionaries, to use both vernacular and Latin names when referring to or describing species, regarding these names as interchangeable in the texts.Latreille (1804:252), in his redescription of Parandra, used the vernacular name "La Parandre lisse", in conjunction with the Latin name "parandra laevis" (as "lisse" was the vernacular name given by DeGeer (1774) for Attelabus glaber).Th e name "laevis" is clearly being used for A. glaber DeGeer, with the reference to the description and fi gure of that species in DeGeer (1774) being given in the text.Latreille (1804) clearly states that the genus is being erected for one species.Th e name "laevis" in this text is not being used as a replacement name in the sense we would understand it today, but under the ICZN (1999) it would be regarded as such.In 1806 Latreille published a fi gure that he believed to be the male of A. glaber DeGeer.Th e text relating to the plates was published in Latreille (1807).In this text he again uses both the vernacular name "Parandre lisse" in conjunction with the Latin name Parandra laevis.However in the legend to the plate only the name Parandra laevis is used.
In the 1807 text Latreille synonymises Tenebrio brunneus Fabricius, 1798 and Tenebrio purpurascens Herbst, 1799 (now Neandra Lameere, 1912) under "Parandra laevis".It is therefore clear from the texts of Latreille (1802Latreille ( , 1804Latreille ( , 1807) ) and Olivier (1803) that Latreille believed that the genus Parandra contained only one species, the "Parandre lisse", Attelabus glaber of DeGeer.It is not until Schönherr (1817) that the name Parandra laevis is used to indicate a species distinct from Attelabus glaber DeGeer.Gyllenhal (1817:145) in the appendix of Schönherr's Synonymian Insectorum (Descriptions Novarum Specierum) states that he has seen DeGeer's "type" (at that time, apparently, the male(s) type(s) was(were) already lost: "Exemplar unicum femininum") of Attelabus glaber and that Latreille's 1806 fi gure, "Parandra laevis", represented a different species.He uses the name "Parandra laevi" (sic) for this species presumably be-cause it is the name used in the fi gure legend of Latreille (1806) and that it was politic at that time to attribute the name of a species to the person who "discovered" it rather than the reviser.Th is is refl ected in Schönherr's (1817:334) main text where Parandra glabra is attributed to DeGeer, Parandra laevis to Latreille, and Parandra brunnea and Parandra purpurea listed as distinct species.Unfortunately, by using the same Latin name (Parandra laevis), that Latreille used for "Parandre lisse" (A. glaber), for the species represented in the 1806 fi gure which Schönherr / Gyllenhal believed to be distinct from Attelabus glaber DeGeer, Schönherr (op.cit.) created a junior homonym.
Latreille (1818: 523) acknowledges and accepts Schönherr's (Gyllenhal's) view that Attelabus glabra DeGeer, 1774 and the 1806 fi gure of "Parandra laevis" are diff erent species.He comments on the distribution of the two species and morphological differences of the specimens he has in his possession from the Antilles and Brazil.He particularly notes that the mandibles of the male in P. glabra are strongly curved and bidentate at the apex with a small tooth on the inner edge whereas in "laevis" the inner edge in the male is bidentate and the apex simple or weakly forked (however it is not clear if he was referring to either the inner element of the apical trifurcation as a second inner tooth, the basal tooth if the mandibles were open or a minor male).He disagrees that Parandra brunnea is a good species.
Th e form of the mandibles would be the "key" for application of the Article 70.3 of the ICZN (1999), for selecting a new type species for Parandra.Unfortunately the description of Latreille ( 1802) is not precise enough to determine if he was describing the mandibles of Parandra laevis sensu auctorum, or the mandibles of Attelabus glaber DeGeer.
Latreille (1818) published a further fi gure of Parandra laevis in the plates of the Encyclopédie Méthodique with some diff erences from the 1806 fi gure, principally the position of the jaws (fi gure legend p. 19 spelt Perandra corrected to Parandra p. 39).Th e text relating to the plates was not published until 1825.In this volume the senior author is Latreille but the text on Parandra was written by LePeletier and Audinet-Serville.In the 1825 text "Parandre glabre; Parandra glabra" is attributed to Schönherr (1817) and "Parandre lisse; Parandra laevis" to Latreille (1807).Th us the connection between "lisse" = glaber of DeGeer = laevis was lost.
Gyllenhal in Schönherr (1817) distinguished P. laevis Latreille as a separate species from P. glabra (DeGeer) with reference to the Latreille 1806 fi gure and 1807 text, but there is no mention in Gyllenhal's text of any actual specimens or that he had seen Latreille's material.Research has failed to locate any specimens determined as Parandra laevis Latreille in the Gyllenhal collection although there is a major male specimen of Parandra laevis in the Schönherr collection from Haiti.Annotations in Schönherr's personal copy of Synonymia Insectorum (1817) in the NRS indicates that he had this specimen in his lifetime (Julio Ferrer pers.comm.) and it probably represents his and Gyllenhal's species concept of Parandra laevis Latreille 1806, 1807.
Post 1818 all subsequent authors when dealing with Parandra have regarded Parandra glabra (DeGeer) and P. laevis Latreille (1804Latreille ( , 1807) ) as separate, valid species.All failed to realize that prior to 1817 the name Parandra laevis as used by Latreille did not defi ne or constitute a separate species from Attelabus glaber DeGeer.
We know that Dejean acquired the Latreille cerambycid collection after his death in 1833.In his 1837 catalogue Dejean attributes the female of Parandra laevis Latreille to another species Parandra cayenensis Dejean, now regarded as Parandra punctatissima Th omson, 1861 and placed in Acutandra Santos-Silva, 2002.Dejean did not use the name Parandra glabra (DeGeer) which may be why Kirby (1837:166) erected a new genus Gnathophorus for Attelabus glaber DeGeer, not realizing that it had already been placed in the genus Parandra by Latreille, 1802. Westwood (1839:358), in a footnote, refers to Attelabus glaber DeGeer as the type species of Parandra and mentions that Kirby (op. cit.) proposes to call it Gnathophorus but fails to comment on the fact that this would makes Kirby's name a junior synonym of Parandra.
Th e Dejean catalogues were very infl uential in their time and many subsequent synonyms and taxonomic problems can be attributed to them.Th omson (1867) validated many of Dejean's Parandra species names hence the large number of synonyms currently listed.
Chevrolat acquired the Dejean cerambycid collection after 1845.Chevrolat (1862:275) states that he has Latreille's "type" of Parandra laevis.Th e Chevrolat collection was divided and sold before his death with the cerambycids coming to the BM(NH) via J. Bowring in 1863.In the manuscript catalogue of the Chevrolat collection in the BM(NH), four specimens are listed as being determined as Parandra laevis Latreille (both sexes).Unfortunately none of the specimens bear Latreille's original determination labels having been relabelled by Dejean.Only one male is labelled as "laevis" Latr.from Latreille's collection.It bears an original handwritten locality label "St.Domingae" (Santo Domingo), the locality given by Th omson (1864:316) in his type species designation and by Lameere (1902:86).Th e other specimens that have been identifi ed as coming from Latreille's collection (ex.Dejean collection), were given a variety of diff erent names by Dejean, these are all now determined as Neandra brunnea (Fab.)confi rming the fact that Latreille's 1807, 1818 concept of Parandra laevis, was a composite one.Th e Parandra laevis male from the Chevrolat (ex.Latreille collection) was labelled as "type", by C. O. Waterhouse, a former BM(NH) curator, with reference to Chevrolat (1862), shortly after it was acquired in 1863.It is thought that the BM(NH) red type label was probably added by K. G. Blair (former curator) ca.1930s.Since 1863 this specimen would have been regarded as the "type" of Parandra laevis Latreille, 1804 by subsequent authors and maybe regarded as representing the species concept of P. laevis Latreille as it is accepted today.
As explained above, the name Parandra laevis as used by Latreille (1804Latreille ( , 1807) ) does not represent a species separate from P. glabra DeGeer, 1774.Th e fi rst author to establish Parandra laevis as being a diff erent species from Parandra glabra was Schönherr (1817).
Th e fi gure given by Latreille (1806), that Schönherr and Gyllenhal (1817) cited in their text as representing a second species to Parandra glabra and the subsequent 1818 fi gure of Latreille are too crude to be sure of the species they represent.We know from the Latreille 1807, 1818 texts that his species concept of "laevis" was a composite one.To resolve any confusion as to the identity of the species currently known as Parandra laevis, Latreille 1804 a new description, and name is provided (to resolve homonymy) and a lectotype selected.It has been decided to designate the specimen labelled as the "type" of Parandra laevis Latreille from the Latreille collection cited by Chevrolat, 1862, now in the NHM(UK), as the lectotype of Parandra laevis Schönherr, 1817.It is not possible to determine, without doubt, which Latreille's specimen is the holotype of P. laevis Schönherr, 1817, and the specimen fi gured by Latreille (1806), (ICZN 1999: Articles 73.1.4, 72.4.2, 75.1).Latreille 1818 states that the 1806 fi gure was drawn from a male from the Antilles.Selecting this specimen, which is from Santo Domingo and which has been cited in the past literature as the "type" of Parandra laevis Latreille, 1804 (see above), maintains the taxonomic stability and species concept of Parandra laevis Latreille, 1804 as it is know and accepted today (ICZN 1999: Article 60.3, 72.4.1).
General colouration of integument a vitreous light chestnut to deep-reddish brown, head may be darker in colour than rest of dorsum, particularly in male; apices and inner margins of mandibles, anterior margin of head, post ocular area, margins of pronotum, scutellum, elytral suture, episternum and anterior margins of femora normally dark to pitchy brown.
Male (Figs 1,3).Width of head (Figs 5,6) plus eyes equal to that of pronotum at anterior angles; length of mandible equal to that of head, fi nely and densely punctured, punctures slightly smaller than those on disc of head, becoming slightly coarser at apices; apices of mandible trifurcate; mandible in major male (Fig. 5) robust with a large tooth on inner edge just above the middle (see Santos-Silva 2002:36), size of sub-median tooth more or less equal to that of dorsal inner apical tooth which is in turn slightly shorter than terminal element but longer and broader than ventral apical tooth, ventral tooth (not visible when viewed dorsally) small and somewhat indistinct, apical teeth may be worn or broken; mandible in minor male (Fig. 6) less robust, submedian inner tooth broadly fl attened, width twice that of inner apical tooth, outer margin of sub-median tooth weekly emarginated; there is also a large basal tooth which is similar in size to sub-median tooth but this is only visible if the jaws are widely open; mandibular dorsal carina distinct, elevated from the base of mandible to just beyond sub-median inner tooth, inner edge of mandible with a fringe of long, pale, setae which project beyond margin and over sub-median tooth.Dorsal surface of the head (Figs 5,6) more or less evenly convex with a weak median longitudinal sulcus, disc moderately and fi nely punctured, punctures irregular, size and distribution similar to that on disc of pronotum, becoming distinctly larger and coarser towards the ocular carina and occiput, post occipital area and genae coarsely and abundantly punctuate, size of punctures at least three times that of largest on pronotum (maybe confl uent in places), dense punctuation extending beneath head to gula which is medially impunctate; dorsal ocular carina distinct, moderately elevated, extending from mid level of eye to base of jaw, width more or less equal to the base of mandibular carina.Frons weakly depressed at sides, prior to anterior margin, resulting in a slight median, transverse ridge at level of anterior margin of antennal insertions.Clypeus weakly elevated medially with a strong median projection; projection at least three times wider than long; anterior margin weakly emarginated.Eye (Fig 8) very weakly emarginated; posterior ocular edge very distinct; length of eye twice that of median width, when viewed laterally; ventral ocular lobe only slightly broader than dorsal lobe.Sub-mentum with similar strong, dense punctuation, elevation of sub-mentum more of less equal to that of genae, delimited behind by a shallow transverse sulcus, punctures with a long pale seta which is at least fi ve times the length of the puncture.Carina of ventral sensorial area of the antennomeres III-XI (Figs 8, 9) distinctly elevated, visible from the side; pilosity of the antennomeres III-XI pale, distinct, longest setae forming posterior fringe on segments, setal length almost equal to that of segment.
Pronotum (Figs 5,6) transverse, somewhat fl attened dorsally, maximum width at anterior angles, being slightly greater than that of elytra at humeri, anterior angles rounded not strongly projected; anterior margin very weakly concave; posterior margin weakly sinuate; lateral margins narrow but well defi ned, becoming gradually but distinctly concave after anterior third with minimum width just prior to posterior angles  that are weakly projected; disc fi nely and irregularly punctuate, punctures fi ner than those on disc of head, becoming gradually larger towards margins, largest punctures not greater than those on anterior area of head.Elytron 3.8 times longer than broad, maximum width at humeri, becoming only slightly narrowed just prior to apex which is broadly rounded; surface adjacent to median suture fi nely punctate, punctures equal in size to smallest on disc of pronotum, distribution irregular, punctures on average 4-5 times their own width apart becoming slightly larger and denser sub-laterally below humeral third.Metepisternum distinctly and irregularly punctuate, punctures of similar size or slightly smaller than those on adjacent area of metasternum, becoming denser near anterior margin; punctures on metasternum large and distinct adjacent to metepisternum 2.3 times their own width apart, becoming distinctly fi ner towards metasternal suture.Apical ventrite of abdomen only slightly longer than preceeding segment, more distinctly punctured than rest of abdomen with long pale sub-erect setae.Femora minutely and indistinctly punctured throughout, setae minute and indistinct.Tibiae becoming strongly dilated towards apex, width at apical margin at least 4× that at base, punctures minute, as on femora, setae minute just extending beyond margins of punctures, ventral fringe short; protibiae somewhat fl attened giving narrow, sharp dorsal edge.Metatarsomere V approximately as long as I-III combined.Body length (including mandibles) 10.3-18.7 mm; prothorax: length 2.2-3.3 mm; anterior width, 3.0-4.8mm; posterior width, 2.5-4.0 mm; elytral length, 6.2-11.5 mm.
Female (Fig. 3).Head less robust, width plus eyes slightly narrower than that of pronotum at anterior angles.Eyes slightly broader and less pronounced than in male.Mandible (Fig. 7) subtriangular, length equal to two thirds that of head, more coarsely punctured than in male, punctures on dorsal surface irregular, largest equal in size to post ocular punctures; dorsal carina present but less well defi ned than in male; ventral apical tooth small (not visible dorsally); dorsal inner edge with small rounded preapical tooth, basal half with a broad fl attened, weakly sinuate tooth which is slightly more pronounced on left mandible.Clypeal projection not broader than long, distinctly narrowing towards apex.Pronotum (Fig. 7) similar to male but with anterior two thirds slightly more rounded at sides prior to becoming narrowed and with anterior angles less pronounced.Apical ventrite of abdomen twice length of preceeding segment, puncturation slightly denser than in male and setae shorter.

Lectotype
As explained above, we have chosen the specimen from Latreille's collection, deposited at NHM(UK), as the Lectotype of Birandra (Yvesandra) latreillei.Th is specimen has the following labels (Fig. becomes a junior synonym of Parandra Latreille, 1802 and all the species currently standing as Parandra (Parandra) Latreille, 1804 in Monné (op.cit.) now require a new generic name.Th e fi rst available name is Birandra Santos-Silva, 2002 (Bousquet 2008) (see note on Protospondylis below).Birandra is currently a subgenus of Parandra Latreille (Monné op.cit.), thus the species currently standing as Parandra (Birandra) (Monné op.cit.) will become Birandra (Birandra) Santos-Silva, 2002.Th e species standing as Parandra (Parandra) Latreille (sensu Santos-Silva 2002) will become Birandra (Yvesandra) nom.n.Although the nomenclatural problems regarding Parandra and P. laevis were known to us before Bousquet (op.cit.) published his work, he was the fi rst to point out these problems.Th us, we chose the name Yvesandra (feminine gender) to honor Yves Bousquet.