A new species of Speonemadus from Portugal, with the revision of the escalerai-group (Coleoptera, Leiodidae) Author Reboleira, Ana Sofa P. S. Author Fresneda, Javier Author Salgado, José Maria text European Journal of Taxonomy 2017 2017-01-19 261 1 23 journal article 32199 10.5852/ejt.2017.261 708e152e-4450-4439-84a9-3c56148cd082 885318 C59239D1-0BF0-446A-AF6E-71234D01B23D Speonemadus algarvensis sp. nov. urn:lsid:zoobank.org:act:5CFDCE6C-71CE-4763-89BD-B8A68E945A80 Figs 1 , 6–7 , 16 , 21 , 26–31 Diagnosis A species of Speonemadus with a total length of 4.0– 4.9 mm , belonging to the escalerai -group, and with typical features of the group; characterised by having the 2nd, 4th, 5th and 7th antennomeres equal or subequal in length; a slightly transverse and hexagonal pronotum; and a long, raised and elevated protibial keel in males, with the apex moderately sharp. Etymology The species epithet is derived from the type locality, the southernmost province of Portugal, the Algarve. Material examined Holotype PORTUGAL : ♁, Algarve , Loulé , Gruta do Vale Telheiro , coordinates: WGS 84: 37º10′14″ N , 008º02′06″ W , 29 Dec. 2009 , S. Reboleira leg. ( ZMUC number 00036257). Paratypes PORTUGAL : 38 ♁♁ and 30 ♀♀ , same data as holotype; 11 ♁♁, 18 ♀♀ , same locality as holotype, 13 Mar. 2009 , S. Reboleira leg. ; 53 ♁♁, 63 ♀♀, 24 May 2009 , S. Reboleira leg.; 9 ♁♁, 9 ♀♀ , Algarve , São Brás de Alportel , Algarão do Remexido , 15 Mar. 2009 , S. Reboleira leg. ; 1 ♀ , same locality, 23 May 2009 , S. Reboleira leg. ; 31 ♁♁, 27 ♀♀ , same locality, 25 May 2009 ; S. Reboleira leg. ; 1 pupa , 1 ♀ , same locality, 5 Sept. 2009 , S. Reboleira leg. ; 3 ♁♁, 3 ♀♀ , same locality, 29 Dec. 2009 , S. Reboleira leg. ; 4 ♀♀ , Algarve , Moncarapacho , Gruta da Senhora , 29 Dec. 2009 , S. Reboleira leg.; 1 ♁ , same locality, 24 May 2009 , S. Reboleira leg. (SR, CASR , CJMS , CZULE , CFL , CPB , ZMUC ). Figs 1–5. Speonemadus of the escalera i- group habitus. 1 . S. algarvensis sp. nov. (Gruta do Vale Telheiro, Portugal). 2 . S. angusticollis ( Kraatz, 1870 ) (Rute, Spain). 3. S. bolivari ( Jeannel, 1922 ) (Ardales, Spain). 4 . S. breuili ( Jeannel, 1922 ) (Motillas, Spain). 5 . S. escalerai ( Uhagón, 1898 ) (Jumilla, Spain). Scale bar = 1 mm. Table 1. Mean measurements (in millimeters) of the antennomers of Speonemadus of the escalerai - group (L) length; (W) width.

1 st	2 nd	3 rd	4 th	5 th	6 th	7 th	8 th	9 th	10 th	11 th
Speonemadus algarvensis sp. nov.
holotype ♁	L	0.189	0.195	0.212	0.196	0.194	0.176	0.196	0.117	0.166	0.137	0.221
W	0.065	0.056	0.056	0.056	0.056	0.056	0.075	0.061	0.088	0.088	0.090
paratypes ♁	L	0.191	0.195	0.208	0.194	0.192	0.170	0.194	0.116	0.163	0.138	0.220
W	0.065	0.056	0.055	0.055	0.055	0.055	0.074	0.060	0.082	0.082	0.082
paratypes ♀	L	0.185	0.195	0.202	0.181	0.179	0.161	0.179	0.105	0.140	0.122	0.225
W	0.066	0.057	0.055	0.053	0.053	0.059	0.069	0.064	0.088	0.088	0.092
Speonemadus angusticollis ( Kraatz, 1870 )
♁	L	0.182	0.208	0.226	0.195	0.200	0.153	0.192	0.114	0.161	0.143	0.231
W	0.078	0.066	0.069	0.065	0.068	0.068	0.087	0.078	0.104	0.104	0.107
♀	L	0.177	0.195	0.205	0.191	0.195	0.150	0.190	0.107	0.159	0.143	0.231
W	0.078	0.068	0.065	0.065	0.068	0.065	0.085	0.073	0.091	0.091	0.094
Speonemadus breuili ( Jeannel, 1922 )
♁	L	0.174	0.177	0.217	0.186	0.186	0.170	0.186	0.117	0.163	0.140	0.222
W	0.078	0.059	0.059	0.056	0.056	0.060	0.072	0.061	0.085	0.088	0.088
♀	L	0.194	0.199	0.228	0.196	0.198	0.179	0.198	0.120	0.155	0.135	0.237
W	0.074	0.060	0.059	0.057	0.057	0.057	0.072	0.065	0.096	0.100	0.100
Speonemadus bolivari ( Jeannel, 1922 )
♁	L	0.182	0.192	0.228	0.192	0.203	0.189	0.205	0.117	0.156	0.135	0.231
W	0.066	0.059	0.055	0.055	0.060	0.062	0.073	0.064	0.088	0.088	0.094
♀	L	0.177	0.182	0.198	0.169	0.190	0.160	0.191	0.107	0.153	0.134	0.228
W	0.068	0.061	0.056	0.056	0.059	0.061	0.072	0.061	0.083	0.083	0.086
Speonemadus escalerai ( Uhagón, 1898 )
♁	L	0.190	0.194	0.217	0.196	0.195	0.160	0.195	0.114	0.176	0.157	0.246
W	0.075	0.060	0.062	0.062	0.062	0.065	0.078	0.074	0.101	0.104	0.104
♀	L	0.200	0.207	0.230	0.208	0.208	0.169	0.207	0.120	0.172	0.156	0.248
W	0.078	0.072	0.073	0.074	0.074	0.077	0.099	0.082	0.109	0.111	0.111

Description BODY. Length 4.03–4.90 mm (males) and 4.00– 4.89 mm (females) width 1.46–1.58 mm (males) and 1.44–1.60 mm (females) ( Figs 1 , 26–27 ). Color reddish brown, darker in central area of head and lighter on antennae and legs. Integument evenly covered with a yellowish pubescence, thin, short and fattened, slightly raised in forehead area and clypeus. Head retractable; head sculpture composed of a tight mesh of small dots. Eyes developed with pigmented ommatidia ( Fig. 27 ); antennae long ( 1.9 mm ) and reaching almost one third of basal area of elytra; antennomeres ( Figs 26–27, 29 ) slender, all clearly longer than wide; 2th, 4th, 5th and 7th antennomeres equal, 3rd slightly longer, 6th shorter and 8th almost twice as long as wide ( Table 1 ); tip of antenna with sensorial organ ( Fig. 29 ). PRONOTUM ( Figs 6–7 ). Slightly transverse, maximum width/length ratio = 1.29–1.43 (males) and 1.28– 1.38 (females), with maximum width at middle; almost fattened; basal impressions not observed; lateral and basal margins weakly beaded; pronotal sides regularly curved anteriorly, posteriorly almost straight and converging posteriorly; basal angles obtuse with apex almost rounded. Pronotal base narrower than base of elytra; middle area gently arched, lateral areas almost straight. Pronotum surface uniformly granulate, well defned laterally and more indistinct in disk area. ELYTRA. Elliptical, very elongate and convex, except along suture where they are fattened in the central area; maximum length/width ratio = 1.85–1.98 in males and 1.95–2.02 in females; elytron apex rounded; transverse striations visible, clearly separated, shallow and perpendicular to elytral suture ( Fig. 28 ); sutural groove present and well marked along entire elytron, sub-parallel to suture and clearly convergent. Metathoracic wings fully developed. LEGS. Relatively long and thin; protibial keel ( Fig. 16 ), which is high and long, elevated distally, with a well marked vertex, sharp at apex; frst three protarsomeres dilated ( Fig. 30 ) in males, frst slightly narrower than apical area of protibia (tarsus/tibia ratio = 0.86); mesotibia curved, metatibia straight. Figs 6–15. Speonemadus of the escalerai -group pronotum. 6–7 . S. algarvensis sp. nov. (Gruta do Vale Telheiro, Portugal), respectively ♁ and ♀. 8–9 . S. angusticollis ( Kraatz, 1870 ) (Rute, Spain) respectively ♁ and ♀. 10–11 . S. bolivari ( Jeannel, 1922 ) (Ardales, Spain) respectively ♁ and ♀. 12–13 . S. breuili ( Jeannel, 1922 ) (Motillas, Spain) respectively ♁ and ♀. 14–15 . S. escalerai ( Uhagón, 1898 ) (Jumilla, Spain) respectively ♁ and ♀. Scale bars = 1 mm. Male Some variability is observed in male paratypes ( Table 1 ). Genital segment elongate, 1.5 × longer than wide; composed of a tergum and two pleurites, all with small setae in apical area; sternum reduced to a long, narrow longitudinal piece, sharp apically. Aedeagus length = 1.2 mm (1.3 if parameres are included) ( Fig. 21 ). Median lobe, lanceolate in dorsal view, with elongate and curved apex close to dorsal face; sharp apex with marked shoulders at margins; basal lamina developed, almost two times shorter than middle lobe; ventral blade of tegmen short and poorly defned. Parameres, in dorsal view, sub rectilinear, slightly curved medially on inner side, with apical areas rounded and slightly truncated on inner side; four thin setae at apex, three subequal upper setae and a larger one, close to largest setae, another seta, shorter, more robust, tooth-like and highly sclerotized. Inner sac with two long rows of spinules surrounding two long parallel rows of sclerotized teeth arranged in a zipper-shape ( Figs 21 , 31 ). Female Generally smaller, with a less transverse pronotum ( Fig. 7 ) and longer elytra, but same body width. No sexual dimorphism observed in colour, pubescence, sculpture, granularity, striation, mesotibia or metatibia. Females have a slender protarsus; antennomeres proportionally thicker and elytral apex notched and serrate. Seventh uroventrite with a narrow slit in middle of posterior margin and 8th uroventrite with posterior margin as a soft membranous arc, its ventral spine is short, wide and slightly sharp at apex. Ratio of frst tarsus to apical part of tibia: 0.84–0.89. Female genitalia typical of genus as described by Giachino & Vailati (1993) and Salgado et al . ( 2008 ) . Figs 16–20. Speonemadus of the escalerai -group male protibia. 16 . Speonemadus algarvensis sp. nov. (Gruta do Vale Telheiro, Portugal). 17 . S. angusticollis ( Kraatz, 1870 ) (Rute, Spain). 18 . S. bolivari ( Jeannel, 1922 ) (Ardales, Spain). 19 . S. breuili ( Jeannel, 1922 ) (Motillas, Spain). 20 . S. escalerai ( Uhagón, 1898 ) (Jumilla, Spain). Scale bar = 0.5 mm. Figs 21–25. Speonemadus of the escalerai -group, aedeagus. 21 . Paratype of S. algarvensis sp. nov. (Gruta do Vale Telheiro). 22 . S. angusticollis ( Kraatz, 1870 ) (Rute) . 23 . S. bolivari ( Jeannel, 1922 ) (Ardales) . 24 . S. breuili ( Jeannel, 1922 ) (Motillas) . 25 . S. escalerai ( Uhagón, 1898 ) (Jumilla) . Scale bar = 0.5 mm. Affnities The morphological differences that separate Speonemadus algarvensis sp. nov. from all other species of the S. escalerai -group are pointed out in the key for the group. Morphologically, the most closely related species are Speonemadus bolivari and S. breuili , which are also the closest geographically ( Fig. 32 ). However, S. algarvensis sp. nov. can easily be distinguished from S. bolivari by the different shape of the male protibial keel, by the antennomeres ratio and the shape of the aedeagus, with the median lobe apically more narrow and elongate. Differences can also be observed in the females of S. algarvensis sp. nov. , with the median posterior slit of the 7th uroventrite slightly narrower and the apex of the ventral spine of the 8th uroventrite less sharpened. The differences regarding S. breuili are much more clear in the shape of the protibial keel. Figs 26-31. Speonemadus algarvensis sp. nov. scanning electron micrograph. 26 . Dorsal view with Laboulbeniales Stichomyces conosomatis pointed out. 27 . Ventral view, with detail of phoretic acari. 28 . Details of elytron stria and setation. 29 . Tip of the antenna. 30 . Male protarsus with detail of the ventral pads of the frst tarsomere. 31 . Posterior dorsal view of the evaginated aedeagus. Scale bars 26–27 = 100 μm, 28–31 = 10 μm. There is a strong geographic isolation between karst areas colonized by the previous three species, which have the most southern distribution of the group ( Fig. 32 ). The male specimen collected on 4 Jan. 1940 , in the cave Igrejinha da Soídos, located in Alte, municipality of Loulé ( Jeannel 1941 ), deposited in the Natural History Museum of Paris (MNHN), has been examined and belongs to the new species S. algarvensis sp. nov. , it had previously been identifed as S. angusticollis . The remnants of another specimen collected in the cave “Berrocal do Esguincho”, also in the Loulé municipality, on 6 Dec. 1983 and cited by Blas ( 1985 , 1989 ), are also included in this new species. Both aforementioned caves are included in the same karst area as the type localities of S. algarvensis sp. nov. , where only this species of Speonemadus has consistently been collected. The specimens reported from the Algarve massif as Speonemadus angusticollis by Jeannel (1941) , Blas (1985 , 1989 ), Reboleira et al. (2010 a , 2010b, 2011b , 2012a ) and Reboleira (2012) should now be included in S. algarvensis sp. nov. Biology and ecology Despite lacking some of the typical traits of cave-adapted species, i.e., eyes, severe depigmentation and extreme body and appendages elongation, Speonemadus algarvensis sp. nov. was never found at the surface, leading us to classify it as a subterranean species. It has been sampled all year round and it has never been collected at the surface in the areas surrounding the sampled caves. The largest population was found in Vale Telheiro Cave, which matches with the general high biodiversity pattern for other groups of troglobionts in Portugal ( Reboleira 2012 ; Reboleira et al. 2015 ). Some specimens of S. algarvensis sp. nov. have the ectoparasitic fungus Stichomyces conosomatis Thaxt., 1901 (of the order Laboulbeniales ) attached to the cuticle (Santamaria pers. com.) ( Fig. 26 ); this represents the frst record of the fungal species for the Portuguese territory. This fungus species has only been previously found in Staphylinidae beetles of the genus Sepedophilus Gistel, 1856 ( Haelewaters et al. 2015 ), that was also found in these caves (Reboleira unpublished). The remarkable discovery of this Laboulbeniales species on a new host of the family Leiodidae is the frst case of host shifting following an ecological opportunity ( sensu De Kesel & Haelewaters 2014 ), in the subterranean environment. Also phoretic undetermined mites were observed on some specimens ( Fig. 27 ). Speonemadus algarvensis sp. nov. shares its habitat with highly subterranean-adapted species such as the detritivorous woodlice Cordioniscus lusitanicus Reboleira & Taiti, 2015 and Trogleluma machadoi (Vandel, 1946) ; the millipedes Acipes machadoi Enghoff & Reboleira, 2013 , A. biflum Enghoff & Reboleira, 2013 , Boreviulisoma barrocalense Reboleira & Enghoff, 2013 and a new species of Archipolydemus Attems, 1898; the campodeid Litocampa mendesi Sendra & Reboleira, 2010 ; the nicoletiid Squamatinia algharbica Mendes & Reboleira, 2012 ; and some predators: the pseudoscorpions Chthonius minutus Vachon, 1940 ; Titanobochica magna Zaragoza & Reboleira, 2010 and Lusoblothrus aenigmaticus Zaragoza & Reboleira, 2012 ; the spiders Harpactea stalitoides Ribera, 1993 , Teloleptoneta synthetica (Machado, 1951) and Anapistula ataecina Cardoso & Scharff, 2009 , as well as a species of centipede, Lithobius Leach, 1814 ( Enghoff & Reboleira 2013 ; Reboleira et al. 2010 a , 2010b , 2010c , 2011 a , 2012 a, 2012b , 2013 , 2015 ; Reboleira & Enghoff 2013 , 2014 ). Distribution Speonemadus algarvensis sp. nov. was collected in three caves of the southernmost province of Portugal , the Algarve ( Fig. 32 ), where it seems to be endemic to the central and eastern part of the Algarve karst massif. The species was not found in the most western part of the Algarve massif, where feldwork was also conducted. The karst included in this region is also know as “Barrocal” and is the richest area for subterranean-adapted fauna of the country ( Reboleira 2012 ; Reboleira et al. 2011 a , 2013, 2015 ; Reboleira & Enghoff 2013 , 2014 ).