NEW TAXA OF LAPAROCERUS SCHÖNHERR , 1834 FROM MADEIRA AND PORTO SANTO , WITH A KEY TO THE SPECIES OF THE MADEIRAN ARCHIPELAGO ( COLEOPTERA , CURCULIONIDAE , ENTIMINAE )

As part of a monographic study of the weevil genus Laparocerus Schönherr, 1834, which is very diverse in Macaronesia, this paper proposes some changes to the systematics of the genus and describes a number of new taxa from the archipelago of Madeira. Combining the morphological data with the molecular data presented in a previous contribution, the subgenera Laparocerus and Atlantis are redefined and three new subgenera are described: Atlantodes n. subgen., Pseudatlantis n. subgen. and Wollastonius n. subgen. In addition, six new species from Madeira are described: L. madeirensis n. sp., L. hobbit n. sp., L. serrado n. sp., L. silvaticus n. sp., L. stuebeni n. sp. and L. prainha n. sp., as well as L. cryptus n. sp. from the Ponta de São Lourenço, Porto Santo and Deserta Grande. A key to all known Laparocerus from the archipelago of Madeira is provided.


Introduction
The genus Laparocerus was established by Schönherr in 1834 to harbour two species described by Boheman in the same work, based on material from "Lusitania", a patria that was later clarified as being the island of Madeira.Wollaston, who worked on the Madeiran archipelago fauna in detail (1854,1857,1862,1871), added one more Laparocerus, four species of Omias, the monotypic genus Cyphoscelis Wollaston, 1854, and eighteen species of Atlantis Wollaston, 1854.All these species were ascribed to the genus Laparocerus by Lacordaire (1863), with Cyphoscelis and Atlantis as sections (subgenera) of it.He also created the group "Laparocerides" including several non-Macaronesian genera (Aomus, Elytrodon, Merimnetes, etc.).The related Madeiran genus Lichenophagus Wollaston, 1854, which was placed in another group, has been recently transferred to the Laparocerini Lacordaire, 1863 by Alonso-Zarazaga & Lyal (1999).
Since the initial studies of Wollaston there have been few additions to the list of Madeiran species (Uyttenboogaart, 1940;Roudier, 1958Roudier, , 1961Roudier, , 1963)), but the number of Laparocerus and Lichenophagus species increased considerably due to further studies by Wollaston and other authors mainly in the Canary Islands (with several subgenera being established).The presently known distribution of the genus Laparocerus covers the archipelagos of Madeira, Selvagens and the Canaries, with a single species in mainland Africa (West Morocco).
This paper is part of a comprehensive systematic revision of the genus Laparocerus.A study of the type material of all species names ascribed to the genus yielded 24 valid species for the Madeiran archipelago (Machado, 2006).Preliminary results from a molecular analysis (Machado et al., 2008) suggest that the Madeiran Laparocerus species form a monophyletic group but also that Lichenophagus fritillus Wollaston, 1854 from Porto Santo clusters within the Laparocerus clade.Consequently, Lichenophagus was restricted to Madeiran archipelago species and relegated in status to a subgenus of Laparocerus.Furthermore, Cyphoscelis Wollaston, 1854 was synonymised with the subgenus Laparocerus Schönherr, 1834.A few other changes in the systematics of the genus Laparocerus were suggested in Machado et al. (2008) but not implemented.
The purpose of the present contribution is to describe three new subgenera, seven new species of Laparocerus and to redefine the concept of the sub-genera Laparocerus and Atlantis according to the results of the previous phylogenetic analysis of the Madeiran clade based on molecular data.The systematic scheme here presented is morphologically and molecularly supported and constitutes, as any phylogeny, a mere hypothesis.
A first key to all species from Madeira and Porto Santo is herein provided.Laparocerus piceus Boheman, 1834, the fossil Laparocerus wollastoni Heer, 1857 and Laparocerus obesulus Desbrochers, 1903 are not included as they have already been established as nomina dubia and a synonym, respectively (Machado, 2006).

Materials and methods
This study is based on material collected by the author in the field and borrowed from different institutions and private collections.Unfortunately no specimens of Laparocerus (Lichenophagus) acuminatus (Wollaston, 1854) from Desertas were available for study, and its characters as used in the key were taken from the original description.Holotypes of the new species are deposited in the Natural History Museum of Santa Cruz de Tenerife, Spain.
The names of the collections from which specimens were studied are abbreviated as follows: Photographs were taken with a Nikon digital camera, and drawings were made using a camera lucida attached to a microscope.Measurements were taken with a micrometer fitted to the microscope.The length, width and height of specimens are abbreviated as L, W and H, respectively, where length excludes that of the rostrum and was measured in dorsal view.Eye convexity is expressed as percentage of a theoretical regular ellipsoid or globe protruding from the profile of the head; thus a 50% convexity refers to a hemispherical eye, 10% Graellsia, 64(2), Diciembre 2008, pp.307-328 -ISSN: 0367-5041 to a fairly flat one, and so forth.Abdominal transverse convexity is determined by dividing the maximum height between elytra and ventrites in lateral view by the maximum width of the elytra measured in dorsal view.
The term prorostrum (taken from Damoiseau, 1967) refers here to the anterior and dorsal portion of the rostrum, delimited by the line where the apical declivity usually starts just at the level of insertion of the antennae; the metarostrum is the posterior portion.Spermathecal terminology follows Thompson (1989).
In order to better support the systematic decisions, the present morphological study is here combined with the results of the molecular analysis of Machado et al. (2008), where mitochondrial DNA markers (fragments of cytochrome oxydase subunit II and of ribosomal 16S subunit) were selected as they are standard markers used in many phylogenetic studies (Caterino et al., 2000).
Figure 1 shows the Madeiran clade of Laparocerus according to the Bayesian 50% majority-rule consensus tree obtained from the analysis of mitochondrial cytochrome oxidase II and ribosomal 16S rRNA gene sequence data (a combined dataset of 1023 bp) by Machado et al. (2008, fig.1).The phylogenetic tree has been simplified by removing the Bayesian support values and most Canarian, Azorean and some redundant Madeiran terminals.Subclades representing subgenera have been additionally marked and are consistent with a second tree obtained by these authors (op.cit.fig.2) adding a fragment of the nuclear elongation factor 1-alpha gene for selected taxa.
Full species names, with author and year, are provided in the key.Schönherr, 1834 TYPE SPECIES: Laparocerus morio Boheman, 1834, by original designation.

Subgenus Laparocerus
The monotypic genus Cyphoscelis Wollaston, 1854, erected for C. distorta, was treated by Lacordaire (1863) as a subgenus of Laparocerus.Its peculiar boat-shaped and flattened outline is probably related to a habitat shift (feeding on the ground on dead leaves, e.g. of Euphorbia mellifera), but all its other characters and apomorphies (e.g. the premental setae) are shared with L. morio.Machado et al. (2008) confirmed the close relationship of L. distortus and L. morio (see fig. 1) and thus synonymised Cyphoscelis with Laparocerus, and they further restricted subgenus Laparocerus to a small monophyletic group represented by L. morio, L. distortus and L. chaoensis, with L. cryptus n. sp. to be added According to this new composition, the subgenus Laparocerus is characterised as follows: moderate to large in size (6-13 mm), males larger than females (opposite to the trend in all other subgenera of Laparocerus), scape fine and abruptly capitate (fig.3A), rostrum short and weakly tapering apically (not constricted), prementum with four setae, elytra with vestiture of small appressed scales and additional very short recumbent setae, intermesocoxal process broad and convex; mesotrochanter in male prolonged into a small posterior spine, pro-and mesotibia curved and crenulate on inner face, metatibia in male flattened and expanded (excavated for > 1/2 of its length), aedea-gus with temones (apodemes) sharply demarcated from body, internal sac with two parallel rows of separated stout teeth in basal region.
Based on molecular evidence, Machado et al. (2008) relegated the status of Lichenophagus to that of a subgenus of Laparocerus and restricted its composition to comprise only two Madeiran species, L. fritillus from Porto Santo and L. acuminatus from Deserta Grande.Its diagnostic morphological modifications (body shape and size, form of scape, eye position, scale vestiture) are probably related to a habitat shift, both species living in foliose lichens on exposed rocks, whereas most other Laparocerus species feed on the foliage of shrubs or trees.Wollaston's original description of Lichenophagus is accurate and adequate, but it is noteworthy that males also have a spiculum relictum (sternite VIII) like all other Laparocerus, that the aedeagus is short with a peculiar pre-apical lateral constriction (unsclerotised area) and that the  internal sac is long and tubular with an apically inserted ejaculatory duct (fig.2).
Laparocerus lamellipes, L. calcatrix, L. lauripotens, L. noctivagans and L. vespertinus share a number of characters, namely an elongate shape, clavate scape, additional long elytral hairs and apically expanded male metatibia.They were termed by Wollaston (1854) as "true Atlantides" and form a monophyletic group strongly supported by the molecular analysis conducted by Machado et al. (2008) (fig.1).Laparocerus clavatus, L. lindbergi and L. undulatus have been considered as belonging to the subgenus Laparocerus because of their more capitate scape (figs.3B, D) and the absence of long additional elytral hairs.However, the molecular data clearly relate them to Atlantis and not to Laparocerus (fig.1).In fact, L. undulatus has a typical Atlantis-shaped male metatibia (fig.3G) and in L. clavatus and L. lindbergi (fig.3H) the apical lobe is absent, but the articular area is incipiently or clearly expanded on the internal face of the tibia.This feature of the metatibia seems to be systematically more meaningful, while the shape of the scape and development of additional hairs are apparently subject to homoplasy.
The revised composition of Atlantis here proposed is restricted to a monophyletic group of large species as discussed above, plus L. hobbit n. sp., L. madeirensis n. sp. and L. serrado n. sp., which are related to L. lamellipes, L. vespertinus and L. lindbergi, respectively.Additional characters of Atlantis species are the apically strongly recurved male protibia, the first metatarsal segment being flattened and asymmetrical at its base (except in L. clavatus and L. serrado n. sp.) and the very long internal sac of the aedeagus, with a bifid basal frenum, several areas of denticles and the ejaculatory duct inserted in a ventral pouch (figs.5A, 6A-B).
TYPE SPECIES: Atlantis mendax Wollaston, 1854.Wollaston (1854) termed as "aberrant Atlantides" a group of Atlantis species with a clavate scape (figs.3E-F) and simple tibiae (figs.3K-L) in both sexes, in contrast with the recurved protibia and apically expanded metatibia in the males of the "true Atlantides".In this case, the morphology is not congruent with the molecular data obtained by Machado et al. (2008), which segregate the "aberrant Atlantides" into two distant clusters: (A) L. mendax, L. inconstans, L. instabilis and L. colasi, and (B) L. abditus, L. excelsus and L. schaumii.Group B is genetically more closely related to the "true Atlantides" and Lichenophagus than to group A (see fig. 1).However, in all these species the aedeagus (fig.6C) has a short internal sac with no frenum, and the ejaculatory duct is inserted at the apex (in the middle in a ventral pouch in Atlantis).Based on these relevant internal characters I am reluctant to associate group B with Atlantis and prefer to establish separate subgenera for both groups A and B in order to avoid the obvious paraphyly of the "aberrant Atlantides".
Species of group A, for which the name of Atlantodes is here proposed, are characterised by their elongate shape, absence of rounded scales, clavate scape (faintly curved in basal third or half), densely pilose legs (at least in males), simple tibiae in both sexes and the characteristics of the aedeagus described above.Laparocerus lanatus and L. navicularis (not included in the molecular analysis) as well as L. prainha n. sp. also belong to this subgenus.The size variation within some species (e.g.L. mendax) is remarkable, with some individuals being only half the size of others.

Laparocerus (Laparocerus) chaoensis
The status of species is supported by the molecular data here presented (Table 1 and fig.4).The islet samples studied from Ilheu de Chão, Bugio and the Ilheu do Desembarcadouro (also known as Ilheu da Cevada) in the extreme east of Madeira, cluster together and show a rather high genetic pdistance (ca.8.6-10.8%) of the mtDnA COII with L. morio Nonetheless, the distribution of the species is remarkable (see map on fig.14) so far as Deserta Grande, which is placed between Bugio and Chão, is apparently not inhabited by it, but by L. cryptus n. sp.Roudier (1958) gives a good account of the morphological differences between Laparocerus morio from Madeira proper, and the islet subspecies he described: L. morio vandeli (= L. chaoensis chaoensis) and L. morio cevadae (= L. chaoensis cevadae Roudier, 1958 nov.comb.).Both taxa are well characterised by their aedeagus, being the median lobe bi-sinuate (not straight) in its concave part, and showing a much triangular and shorter apical plate (in dorsal view).The internal sac is armed at base with two parallel rows of 30-32 large denticles (ssp.chaoensis) and 50-70 (ssp.cevadae) instead of the 9-13 of Laparocerus morio or L. cryptus.Moreover, the apical outer angle of the male metatibiae are clearly more protruding than in L. morio, (strongest in L. chaoensis cevadae, fig.3Q) and the normal size of both males and females is generally smaller.
Male specimens from the almost inaccessible rock of Bugio, kindly collected by the park ranger Isamberto Silva (7mm and 6ff, 20-7-2008), bear 27-30 denticles in the aedeagus and show almost the same metatarsal conformation as the males  from Ilheu de Chão.However, their genetic distance (3.8%) is high enough to consider them as a separate subspecies, but we prefer to wait until more material can be examined.In L. chaoensis cevadae, the notch of the internal apical angle of the metatibia is extremely short and preceeded by a sharp point (fig.3Q).Other material examined: More than 150 specimens from Ponta de São Lourenço, Caniçal, Prainha, Porto Santo and Deserta Grande.One pair of this species, confirmed by male genitalia, was collected under a stone at the entrance of the lighthouse in the Ilheu de Chão.An introduction with supplies coming from Deserta Grande is plausible.The islet is abundandtly populated by L. chaoensis, of which more than 100 specimens were collected that same day.
ETYMOLOGY: Adjective derived from the Greek cryptós, hidden, in reference to the veiled identity of this insect (difficult to distinguish morphologically from L. morio. DISTRIBUTION AND ECOLOGY: Laparocerus chaoensis n. sp.occurs in the semi-arid habitats of Ponta de São Lourenço (Madeira) and the islets of Porto Santo and Deserta Grande.It is common in winter and active by night, having been beaten from several plant species (e.g.Atriplex, Cynara, Cistus, Erica, Euphorbia, Galactites, Lotus), and during the daytime it can be easily found under stones.Its vicariant in Madeira proper, L. morio, is more common in summer.
REMARKS: Machado et al. (2008) found that specimens with the habitus of L. morio from Ponta de São Lourenço (East peninsula of Madeira), Porto Santo and Deserta Grande clustered together, showing closer genetic relationship between them (COII p-distance 2.6-3.3% ) than with specimens of L. morio from the rest of Madeira (Table 1).The pdistances obtained with the latter specimens were extraordinarily high (11.0-12.2%);much higher, for instance, than between L. morio and L. distortus (5.9-6.3% which are morphologically very distinct).The hypothesis of L. cryptus being a cryptic species separate from the rest of species in the subgenus is supported by the phylogenetic tree obtained (fig.4), notwithstanding that it is hardly distinguishable from L. morio (see diagnosis).There are no obvious significant allometric differences, except that males of L. chaoensis cryptus usually (but not always) have broader elytra (similar to females).
ETYMOLOGY.The specific epithet refers to the island of Madeira, where the species is endemic, and is an adjective.DISTRIBUTION AND ECOLOGY.Laparocerus madeirensis n. sp. is endemic to the island of Madeira, having being found in the central and eastern forest districts, at high elevation (780-1000 m).It seems to be linked to the hyper-humid laurel forest environment, but, in contrast to most species of the L.lamellipes-group, it is much less common.L. madeirensis n. sp. is active at night and has been beaten from Vaccinium padifolium (Ericaceae), Laurus azorica (Lauraceae) and once from Teline maderensis (Leguminosae).REMARKS.L. madeirensis n. sp.belongs to the group of L. lamellipes within subgenus Atlantis, and is most closely related to L. vespertinus Wollaston, 1854.It is easy to recognise by its elliptical outline (shoulders effaced) and, especially, by the heavily punctured pronotum, which is dull and somewhat uneven in a form unparalleled within the group.The form of the apical anterior lobe of the male protibia varies much in L. noctivagans, but in L. madeirensis it is more constant in shape, subparallel to the axis of the tibia, shortly notched, and never sharp-pointed.This anterior lobe is absent in L. lamellipes, L. hobbit n. sp. and L. vespertinus.DIMENSIONS, holotype (m).Length: total (without rostrum) 9.10 mm, head 2.00 mm, rostrum 0.85 mm, eyes 0.58 mm, scape 2.10 mm, funicle 3.20 mm, articles (1 st / 2 nd / 3 rd / 4 th ) 0.50 /0.64/ 0.38 /0.28 mm, club 0.68 mm, pronotum 2.00 mm, elytra 6.20 mm, and tibiae (pro-/meso-/meta-) 2.65 /2.45 /2.85 mm.Width: head (with eyes) 1.32 mm, head (between eyes) 0.60 mm, rostrum (with pterygia) 1.10 mm, rostrum (minimum dorsal /ventral) 0.60 / 0.90 mm, eyes 0.50 mm, scape (apicad) 0.20 mm, club 0.20 mm, pronotum (anterior /maximum /posterior) 1.45 /2.25 /2.00 mm, and elytra (maximum) 3.55 mm.Height: abdomen 3.00 mm.DIAGNOSIS.Size 8.9-9.1 mm.Similar to L. lamellipes, but scape more sinuous (slightly and uniformly arcuate in L. lamellipes); first article of funicle clearly shorter than second; pronotum laterally more uniformly curved; elytra somewhat narrower and more cylindrical, more depressed at base and suture slightly elevated (keeled) in apical third; additional protruding whitish setae and tufts of thick blackish setae only in apical third of elytra.Legs more robust, with broader tibiae; male protibiae similarly excavated but less curved in middle, with strong brief preapical constriction; 1 st -2 nd tarsal segments incrassated in both sexes, the 2 nd markedly transverse (longitudinal in L. lamellipes); mesotibiae more arcuate.

Laparocerus (Atlantis
ETYMOLOGY.The specific epithet is a noun in apposition which refers to the Hobbits of J. R. R. Tolkien (The Lord of the Rings), a literary fictitious race of people who have big and hairy feet; a metaphor of the swollen and hairy tarsi characteristic of this species.
Graellsia, 64(2), Diciembre 2008, pp.307-328 -ISSN: 0367-5041 DISTRIBUTION AND ECOLOGY.The few known specimens of L. hobbit n. sp. were collected at night in the northern laurel-forest district of Madeira.The specimens from Faja da Nogueira were beaten from Persea indica (Lauraceae) mixed with Laurus novocanariensis (Lauraceae), and the female from São Jorge was beaten from mixed vegetation of Clethra arborea (Clethraceae), Euphorbia mellifera (Euphorbiacea) and Rubus ulmifolius (Rosacea), together with many specimens of L. lamellipes, which is a rather common species.

Laparocerus (Atlantis
ETYMOLOGY.The specific epithet is a noun in apposition taken from the type locality Eira do Serrado, which presumably derives from "serra" = mountain-range, in Portuguese. DISTRIBUTION AND ECOLOGY.The small series of Laparocerus serrado n. sp. was collected on the cliffs adjacent to the old tunnel below Eira do Serrado, in the central mountain range of the island of Madeira.It was beaten at night from Teucrium betonicum (Lamiaceae) growing intermixed with Erica (Ericaceae) and Rubus (Rosaceae).
REMARKS.Laparocerus serrado n. sp. is somewhat intermediate between L. clavatus and L. lindbergi.According to molecular data (fig.1) its sister species is L. lindbergi, but it resembles much more closely L. clavatus, particularly the males, due to the undulated uneven intervals, with bumps clothed with patches of clear scales (this nodosity is much more developed in both sexes in L. clavatus), and by still marked but less prominent (not porrect) humeri in males.The pronotum in L. clavatus is noticeably much wider at the base than anteriorly; in L. serrado the sides are more uniformly and strongly curved, and in L. lindbergi the pronotum is narrower with less curved sides.In the latter species, the male protibiae are extraordinarily recurved and twisted, much more than in any other species of Laparocerus.DNA sequences.GenBank accession numbers: EF583335 (COII) and EF583408 (16S rRNA) from Ribeiro Frio, Madeira.DIMENSIONS, holotype (m).Length: total (without rostrum) 6.05 mm, head 1.30 mm, rostrum 0.62 mm, eyes 0.32 mm, scape 1.48 mm, funicle 1.44 mm, articles (1 st / 2 nd / 3 rd / 4 th ) 0.29 /0.38 /0.18 /0.18 mm, club 0.52 mm, pronotum 1.25 mm, elytra 4.40 mm, and tibiae (pro-/meso-/meta-) 1.74 /1.52 /1.88 mm.Width: head (with eyes) 1.02 mm, head (between eyes) 0.60 mm, rostrum (with pterygia) 0.76 mm, rostrum (minimum dorsal /ventral) 0.60 / 0.70 mm, eyes 0.26 mm, scape (apicad) 0.14 mm, club 0.15 mm, pronotum (anterior /maximum /posterior) 1.17 /1.65 /1.47 mm, and elytra (maximum) 3.15 mm.Height: abdomen 2.40 mm.DIAGNOSIS.Size mm= 5.0-5.9mm, ff= 5,3-6.4mm.Similar to Laparocerus abditus, but integument less shiny, scape as long as pronotum, decumbent pilosity of pronotum and elytra conspicuous but 0.5-0.7xshorter and more compact and uniform (pile 1/4 length of onychium); elytra broader at base and much stouter in profile; tibiae without mucro, protibiae of male almost straight (not shortly incurved at apex).Aedeagus (fig.7B-D), apex in profile sinuous, in dorsal view triangular (straight and blunt in L. abditus).Female tergite VIII tongue-shaped, not triangular (fig.8C).Males and females rather similar in outline and leg configuration (female elytra slightly more quadrangular, with flatter intervals).ETYMOLOGY.The specific name is a Latin adjective meaning wild, derived from silva = forest, used here in reference to the habitat of the species.DISTRIBUTION AND ECOLOGY.Laparocerus silvaticus lives in the moist laurel-forests of the windward part of Madeira, at lofty elevations (800-1350 m).It is common in summer at night and apparently very polyphagous, having been beaten from many different plant species of the forest undergrowth (Plantago, Geranium, Juncus, Hypericum, etc.) as well as from taller bushes like Vaccinium or Erica (Ericaceae).In contrast, L. abditus seems to be restricted to lower altitudes in the dryer leeside of the island, inhabiting scrub vegetation with Euphorbia piscatoria (Euphorbiaceae) or Globularia salicina (Globulariaceae).
ETYMOLOGY.The specific epithet is the name of the type locality Prainha in apposition, which means "little beach" in Portuguese (the insect has preference for sandy soils).DISTRIBUTION AND ECOLOGY.Laparocerus prainha n. sp. is endemic to Madeira.It lives on sandy or mixed sandy-clayish soils in the semiarid Ponta de São Lourenço, feeding on low plants like Lotus glaucus (Leguminosae), but also on ruderals (e.g.Solanum nigrum).There are a few specimens apparently obtained from higher altitudes (Pico do Arieiro and Santo da Serra) but these could well be labelling errors.Its sister species L. mendax lives preferably on sandy soils, in Porto Santo, while L. instabilis selects clayish soils.REMARKS.Laparocerus prainha n. sp.represents in Madeira the phyletic line of L. mendax -L.instabilis from Porto Santo.Because of its moderate size, heavily punctured pronotum and reduced pilosity, it resembles more closely L. instabilis, but DNA sequences (fig. 1) reveal a closer relationship with L. mendax.In all three species, the aedeagus is similarly shaped with an abrupt declivity at the apex of the median lobe in lateral view (fig.6C), but differs in its shape in dorsal view: it is uniformly tapering in L. mendax, preapically swollen in L. instabilis, and shortly constricted in L prainha n. sp.The general pilosity is much more developed in L. mendax, including antennae, dorsum, venter and legs, the first and second segments of funicle are equal, and the puncturation of the pronotum is finer and less dense (smooth surface in between clearly visible).
DISTRIBUTION AND ECOLOGY.Laparocerus stuebeni was found in the south-west end of the island of Madeira (400 m altitude), by shifting debris under a fig tree.The natural habitat of the zone is similar to that inhabited by L. abditus in the eastern part of the island, a dry-scrub vegetation characterised by Euphorbia piscatoria (Euphorbiaceae).