Misolampus goudotii Guerin-Meneville , 1834

Rosas-Ramos, Natalia, Mas-Peinado, Paloma, Gil-Tapetado, Diego, Recuero, Ernesto, Ruiz, Jose L. & Garcia-Paris, Mario, 2020, Catalogue, distribution, taxonomic notes, and conservation of the Western Palearctic endemic hunchback beetles (Tenebrionidae, Misolampus), ZooKeys 963, pp. 81-129 : 81

Misolampus goudotii Guerin-Meneville, 1834

Misolampus goudotii Guérin-Méneville, 1834: 28. Terra typica: “trouvée à Tanger... ... à trois lieues de Tanger, sur les bords d’une rivière, dans le tronc d’un olivier." Vauloger de Beaupré ( 1900), Reitter (1917), Antoine (1949), and Español (1949) among others, wrote the species name with a single final -i. Solier (1848) and Vauloger de Beaupré ( 1900) mentioned the unavailable name: " Misolampus nigrita Dejean", and Español (1954a): " M. moraguesi ".

Misolampus goudotii Erichson in Wagner, 1841: 184 (non Guérin-Méneville 1834). Terra typica: not indicated, but "von Algier" according to Erichson’s (1841) work title.

Misolampus erichsoni Vauloger de Beaupré, 1900: 674 syn. nov. Terra typica: “Algérie: O., Oran...; Daya...; Tlemcen...; Mascara..., Ammi Moussa; A.: Blidah...; La Chiffa...; Margueritte...; forêt de Boghar...; mont Ouarsenis...; forêts de la Grande-Kabylie...".

Misolampus peyerimhoffi Antoine, 1926: 257 syn. nov. Terra typica: "Grand Atlas, région du Glaoui: plateau des Aït Rba...".

Studied material.

Algeria: “Argelia” (Dufour leg.): 1 ex. Morocco - Marrakech-Tensift-Al Hauz: Toufliht, 1483 m, 31°28'34.6"N, 7°26'06.5"W, 11-III-2013: 4 exx. - Meknès - Tafilalet: Ain Leuh, 17-V-1925: 1 ex.; Azrou, 1900 m (Alluaud 79) ( Misolampus goudoti var. laevior Alluaud): 1 ex.; Azrou, 19-V-1925: 1 ex.; Dj. [Yebel] Hebri, 20-V-1925: 1 ex.; Timadit [Timahdite], 21-V-1923: 1 ex. - Tanger - Tétouan: Rif: Beni Siyyel: Bab Ruadi: VI-1932 (C. Bolívar leg.): 6 exx.; Tanger, 1897: 3 exx.; Tanger (M. Escalera leg.) (small square-label pinned): 36 exx., plus 3 exx. only square-labelled; 2 km al O de Bab Berret, 1318 m, 35°00'02.57"N, 4°55'31.91"W, 12-VI-2011: 3 exx.; Crtra. Zinat-Mulay Abdeselam, P-4702, Beni Aros, Yebala, 513 m, 35°22'04"N, 5°32'17"W, 29-IV-2016: 5 exx.; Yebel Bou-Hachem, Beni Aros, Yebala, 1160 m, 35°15'31"N, 5°30'49"W, 12-V-2012: 6 exx.; Crtra. Mulay Abdeselam-Al Hamra, P-4704, Beni Aros, 985 m, 35°15'50"N, 5°25'36"W, 28-XI-2019: 2 exx.; Larache: Yebala: Beni Arós: Yebel Bou-Hachem, 35°15'31"N, 5°30'49"W, 9-VI-2012: 2 exx.; Pinsapar del Talassemtane, 1900 m, 35°08'36.7"N, 5°08'13.0"W, 11-VI-2011: 2 exx.; Bab Taza: Talassemtane, 1401 m, 35°06'10.9"N, 5°08'21.3"W, 27-VII-2013: 1 ex.; Bab Taza: Talassemtane: Plaza de España, 1667 m, 35°09'03.7"N, 5°08'28.6"W, 27-VII-2013: 1 ex.; Casa Forestal, Yebel Lekraa, P.N. Talasemtane, Chefchaouen, 35°07'56"N, 5°08'11"W, 1695 m, 7-VI-2008: 3 exx.; Yebel Talassemtane-vertiente sur, P.N. Talasemtane, Chefchaouen, 35°07'53"N, 5°08'01"W, 1650 m, 11-IV-2011: 4 exx.; P.N. Yebel Tazaot, Pinsapar, P.N. Talasemtane, Chefchaouen, 35°15'N, 5°07'W, 1670 m, 7-V-2011: 2 exx.; Pista hacia Casa Forestal, Yebel Lekraa, P.N. Talasemtane, Chefchaouen, 35°07'45"N, 5°08'09"W, 1700 m, 28-VII-2011: 1 ex.; E. de Yebel Talaousisse, P.N. Talasemtane, Chefchaouen, 35°07'33"N, 5°04'03"W, 1350 m, 1-XII-2018: 2 exx.; Pista hacia Haout Taznout, P.N. Talasemtane, Chefchaouen, 35°08'20"N, 5°07'24"W, 1712 m, 27-IV-2019: 2 exx.; Yebel Tizirhen, Bab Berred, Rif Central, 1585 m, 35°00'54"N, 4°54'57"W, 27-IV-2017: 3 exx.; Yebel Tizirhen, Bab Berred, Rif Central, 1570 m, 35°00'47"N, 4°54'03"W, 28-IV-2018: 1 ex.; Pista de Bab El Kar, Montañas de Fifi, Rif, 1512 m, 34°59'13"N, 5°11'20"W, 2-VI-2019: 2 exx. - Taza - Al Hoceima - Taounate: Iguermalen [Targuist]: Beni Mesdui, VI-1932 (M. Escalera leg.): 6 exx.; Rif: Beni Seddat: Imosiner: VI-1930 (exp. C. Bolívar leg.): 3 exx.; Rif: Beni Seddat: Tizi Taka, VI-1932 (C. Bolívar leg.): 4 exx.; Rif: Beni Seddat: Tizi Taka, VI-1932 (Exp. C. Bolívar leg.): 1 ex.; Rif: Iguermalen (Targuist), VI-1930 (exp. C. Bolívar leg.): 4 exx.; Rif: Ketama, Bab Chiquer, VI-1932 (C. Bolívar leg.): 8 exx.; Rif: Ketama, Bab Chiquer, VI-1932 (M. Escalera leg.): 2 exx.; Rif: Ketama: Tainza, VI-1930 (exp. C. Bolívar leg): 3 exx.; Rif: Ketama: Tidiguin [Tidghine], VI-1930 (exp. C. Bolívar leg.): 1 ex.; Rif: Ketama: Zoco Telata, VI-1932 (M. Escalera leg.): 7 exx.; Lurdeka [?]: 1 ex.; Yebel Tidighin, Azila, Rif central, 1705 m, 34°51'14"N, 4°32'19"W, 29-XI-2019: 1 ex; Carretera P-5420, P.N. Tazzeka, Medio Atlas nororiental, 1000 m, 34°03'N, 4°15'W, 25-XI-2004 (F.J. Martínez leg.): 2 exx. - Souss-Massa-Drâa: Yebel Tual, 28-VII-1934: 1 ex.; Ifni: Yebel Tamarrut [25 km SE Ifni], I-1935 (F. Escalera leg.): 1 ex.; Sidi Ifni: Akarkor, Jbel Toual, 627 m, 29°13'48.9"N, 10°00'44.1"W, 21-I-2020: 4 exx. Spain - Islas Baleares: Mallorca (Mas de Xaxars leg.) ( Misolampus erichsoni ): 2 exx.; Escorca, 26-III-1985, 1 ex.; Escorca, Coll de Femenia, 545 m, 39°51'33.68"N, 2°54'19.27"W, 25-III-2012: 10 exx.; Menorca (Cardona leg.): 2 exx. plus 6 exx. without data; 2 exx.; Menorca: 2 exx.; Algaiarens, 14 m, 40°02'28.3"N, 03°55'28.4"W, 27-IV-2006: 2 exx.

Diagnosis.

With a total length from 10 to 14 mm, this is the largest species of the genus ( Vauloger de Beaupré 1900; Reitter 1917; Antoine 1926; Español 1949). This species is well characterised and isolated within the genus Misolampus by the following traits: fore angles of the prothorax not protruding, almost rounded, forming an obtuse angle at apex; lateral surface of pronotum shallowly rugose, with the rugosity progressively erased towards the dorsal areas that appear smoother, propleural punctation fine and often erased; elytra with longitudinal series of small elongated tubercles, more apparent on the sides of the posterior half of the elytra ( Español 1949, 1954a; Palmer 1998) (Fig. 3A-C ). Female genitalia figured by Palmer (1998). Specimens from the Balearic Islands have been studied karyologically (Juan and Petipierre 1986, 1989; Juan et al. 1993; Pons et al. 1993; Pons 2004), presenting a chromosome number of 20 (2n) ( Juan and Petitpierre 1986, 1989). There is marked geographical variability on the sculpture and shape of pronotum and propleurae, and on the development of elytral tubercles ( Vauloger de Beaupré 1900; Antoine 1949; Español 1954a) (Fig. 3A-C ). Specimens from northern Morocco (excluding the Tingitane Peninsula), Algeria and the Balearic Islands, present a well-developed and evident elytral tuberculation that may form rugose ridges (Fig. 3A ). On the other extreme, elytral tubercles are reduced in the Rif and Atlas populations (Fig. 3B ), to become almost completely absent in the specimens from Sidi Ifni (Fig. 3C ). Pronotum sculpture is formed by fine spaced punctures intermixed with granules, much denser on the sides in the Balearic Islands population (Fig. 3A ); pronotal rugose areas are more marked and extended in the specimens from the High Atlas (Fig. 3B ), and formed by sparse punctation, without granulose areas, in the specimens from Ifni (Fig. 3C ). The anterior edge of the pronotum, in the Rif and Balearic populations, is straight at the middle, while it appears convex in the populations from the High Atlas ( Antoine 1949). The geographic distribution of this variability has been the subject of taxonomic discussion resulting in the proposal of different taxa, here formally synonymised (see synonymic list, and taxonomic discussion).

Geographic distribution.

Distributed throughout Morocco, northern Algeria and the Balearic Islands in Spain ( Antoine 1926, 1949; Español 1949; Löbl et al. 2008). Precise records are well distributed in northern Morocco and Mallorca, scanty in all other areas ( Solier 1848; Lucas 1849; Cardona Orfila 1875; Pérez Arcas 1873; Moragues 1889; Champion 1891; Vauloger de Beaupré 1900; Martínez de la Escalera 1914; Reitter 1917; Peyerimhoff 1919; Antoine 1926; Lindberg 1933; De la Fuente 1934-1935; Palau 1945; Antoine 1949; Español 1949, 1953, 1954a; Cobos 1955; Pardo Alcaide 1955; Kocher 1958; Cobos 1961; Español 1967; Mouna and Arahou 1986; Juan and Petitpierre 1989; Whitehead 1993; París García et al. 2011; Benyahia et al. 2015, 2016; Núñez et al. 2016; Chavanon 2020) (Fig. 4 ). The record from Ceuta, Spain ( Vauloger de Beaupré 1900) corresponds to the mountain Yebel Musa (just 1.5 km west of Ceuta), currently in Moroccan territory (region of Tanger-Tétouan).

The studied materials include recent and old records of populations from the Balearic Islands (Mallorca and Menorca) and from the Moroccan regions of Meknès-Tafilalet, Souss-Massa-Drâa, Tanger-Tétouan, and Taza-Al Hoceima-Taounate. Recent data are available from all four regions, with a large number of localities from the Rif, and less numerous in the Middle and High Atlas. Among these records, we emphasise the re-discovery of the population from the province of Sidi Ifni, in January-2020, 85 years after its original finding, by F. Martínez de la Escalera in 1934 and 1935 ( París García et al. 2011). The latter is a singular population, apparently isolated in the arid mountains near Ifni; its closest known population is located in the Western High Atlas, ca. 250 km to the northeast (Fig. 4A ). The potential distribution map locates high suitable areas for this species along the mountain ranges of northwestern Africa, the coastal and mountain areas in the Tingitane peninsula, and along the coast of Rabat-Salé-Kénitra region. It also identifies areas where the species does not occur as high suitable, including sothwestern Iberia, the Balearic Islands and Sardinia. The Ifni population is located in a very fragmented area of high suitability, suggesting a possible Pleistocene relict status for this population (Fig. 4B ).

Notes on natural history.

Misolampus goudotii is widely distributed over northwestern Africa, though restricted to mountain ranges and adjacent areas: Rif, Middle Atlas, western High Atlas, Beni Snassen mountains, southwestern foothills of the Anti-Atlas (Morocco) and Tellian Atlas (Algerie) (Fig. 4 ). Altitudinal range in the Maghreb from 2 to 2064 m a.s.l., with 70.5% of records above 800 m of altitude (62% above 1000 m). In the Balearic Islands its altitudinal range is lower, between 15 and 718 m a.s.l., but the species is found mainly in areas of mountainous topography (e.g., Serra de Tramuntana in Mallorca). It inhabits a wide range of geological substrates, both acid and basic, from plutonic and metamorphic types to calcareous and dolomitic rocks (see Michard 1976; Vera 2004; Oliveira and Quesada 2019a, 2019b). Misolampus goudotii is a euryecious species that occurs at infra-, thermo-, meso- and supra-Mediterranean thermoclimatic belts, in regions with ombrotypes from arid to hyperhumid ( Benabid 1985; Rivas-Martínez 1987; Le Houerou 1989; Rivas-Martínez et al. 2002; Rivas-Martínez 2007; Sebbar et al. 2013), and occupies a wide variety of forest formations, both coniferous [ Tetraclinis articulata (Vahl) Mast., Abies maroccana Trab., Cedrus atlantica (Endl.) Manetti ex Carrière, Juniperus phoenicea L., Juniperus thurifera L., Pinus nigra J.F. Arnold, Pinus halepensis Mill., P. pinaster ] and broadleaved [deciduous: Quercus canariensis Willd., Quercus afares Pomel, Q. faginea , Q. pyrenaica ; perennial: Quercus ilex , Q. suber , Olea europaea var. sylvestris (Mill.) Lehr] (see Benabid 1982, 1984, 1985; Benabid and Fennane 1994; Bolòs 1997; Charco 1999; Benabid 2000; Taleb and Fennane 2019). It also occurs in areas reforested with pines (pers. obs.) (Fig. 3D, E ). The population of Ifni inhabits mountains (620-1225 m of altitude) at the infra-Mediterranean thermoclimatic belt, probably affected by the proximity to the Atlantic Ocean and consequently by the presence of some degree of cryptic precipitation ( Géhu and Biondi 1998). The vegetation of the area is dominated by open forest of Argania spinosa (L.) Skeels, with sparse cactiform and arbustive Euphorbia L. ( Médail and Quézel 1999; Ruiz and García-París 2015), and large areas covered by formerly cultivated Opuntia Mill (Fig. 3F ).

In the Moroccan Rif, M. goudotii is often encountered under bark, inside fallen logs or stumps, and at the base of dead old oaks (perennial: Q. ilex , Q. suber ; deciduous: Q. canariensis , Q. faginea and Q. pyrenaica ), arbutus trees ( Arbutus unedo L.), wild olive trees ( O. europaea var. sylvestris ), pines ( P. nigra , P. pinaster , P. halepensis ), firs ( Abies maroccana ), and cedars ( Cedrus atlantica ), as already reported partially by Vauloger de Beaupré ( 1900), Cobos (1955, 1961), and Benyahia et al. (2015, 2016). They can also be found under bark of standing dead trees ( A. maroccana , C. atlantica , Q. suber , Q. pyrenaica ). In the Middle and High Atlas, it is usually found under bark and inside large decaying logs of Q. ilex ( Antoine 1926), but also in old decomposing logs of P. nigra and C. atlantica . Mouna and Arahou (1986) collected the species on thuya ( Tetraclinis articulata ) in the Korifla Valley (northwestern Morocco). Sidi Ifni specimens were found within crevices in old dead logs of Argania spinosa , almost buried on the ground of a steep slope (Fig. 3F ). Nearby standing dead trunks were occupied by Nesotes tuberculipennis villarubiai ( Español, 1943) as described by Nabozhenko (2015). In Algeria, they have been found under bark of fallen pines ( Vauloger de Beaupré 1900). In Menorca, it has been found in oak forests of Q. ilex , under bark or under stones and leaf litter ( Cardona Orfila 1875), and in Mallorca it is frequent in decaying wood of fallen pines ( P. pinea ) and old oaks ( Q. ilex ) (Fig. 3D ).

Adult specimens are often found in aggregations. We found aggregations of approximately 15 specimens close together in a single large rotting pine log in Mallorca. We also found aggregations of M. goudotii together with Helops insignis maroccanus (Fairmaire, 1873) ( Tenebrionidae , Helopinae ) under bark of dead trees of Q. suber , A. maroccana and C. atlantica in the Rif Mountains. Whitehead (1993) relates the finding on two occasions of groups of individuals between the annual rings of dead pines ( P. halepensis ) in active colonies of ants of the genus Messor Forel, 1890 and of the species Monomorium bicolor Emery, 1877 (probably another species of Monomorium Mayr, 1855, since the invasive M. bicolor is not present in Balearic Islands; Salata et al. 2019).

Adults are present all year round, but they are more commonly seen in winter and spring in middle and low elevations ( Vauloger de Beaupré 1900; Español 1967; pers. obs.), and in summer at higher altitude ( Antoine 1926), however, Moragues (1889) mentioned collections during the summer in Mallorca.

Figures

Figure 3. Live specimens and habitat of Misolampus goudotii A-C Live adult specimens of Misolampus goudotii from the Balearic Islands (A Cap Formentor, Mallorca) and Morocco (B Toufliht, High Atlas C Akarkor, Sidi Ifni); the specimens selected represent the diversity of sculptural patterns in elytra and pronotum reported for the species D-F A summary of the impressive habitat diversity used by M. goudotii from the Balearic Islands (D Quercus ilex forest at Creu de Menut, Mallorca), to southwestern Morocco (E deep valleys in the Toubkal National Park, High Atlas F Argania spinosa open forests at Jbel Toual in Sidi Ifni). Photographs by MGP and NRR.

Figure 4. A Geographic distribution of Misolampus goudotii. Map for the distribution range of Misolampus goudotii (pale blue spot). Blue dots correspond to the species records, including both recent and old, as well as previously published data. The population from Ifni remains isolated from the main distribution range, by a distance of ca. 250 km B potential geographic distribution of Misolampus goudotii: Red indicates high suitable areas, and blue, areas of low suitability. Species distribution model was generated using MaxEnt v 3.4.1 (Elith et al. 2006) and the set of WorldClim v 2.0 (Fick and Hijmans 2017) environmental variables.