Heriaeus buffoni (Audouin, 1826) is reported for the first time from the Canary Islands, where it was found on Lanzarote. This also represents the first record of the genus in the archipelago. All individuals were collected with pitfall traps installed in nitrophilous synanthropic shrub vegetation near urban areas. Species identification was based on male genitalia only as females were not sampled. A map including all known records from Lanzarote, drawings of the pedipalps and photographs of living and preserved specimens are presented.
The Canary Islands are an archipelago of volcanic origin located off northwest Africa, comprising eight major islands. Lanzarote is the second oldest (15 my) and the easternmost island, being only 140 km off the African coast (Troll & Carracedo 2016). Due to a comparatively low altitude of the island's topography, the humidity of the tradewinds is not retained, which results in little precipitation and desert-like conditions. Thus, together with the high erosion and aridification experienced by this island, Lanzarote exhibits a reduced habitat diversity in comparison with younger central-western islands (Macías-Hernández et al. 2016). Among the more than 500 spider taxa that have been reported for the Canary Islands only 98 have so far been recorded from Lanzarote (Suárez 2018), thus being currently the island with the lowest spider species diversity within the archipelago. However, arachnological studies on this island have been scarce, being mainly limited to taxonomic research focused on specific genera such as Dysdera C. L. Koch, 1837 (Arnedo et al. 2000), Pholcus Walckenaer, 1805 (Dimitrov & Ribera 2007), Loxosceles Heineken & Lowe, 1832 (Planas & Ribera 2015) or the contributions made by Wunderlich (1987, 1992, 2011). Thus, the lack of chorological studies translates into a poor understanding of the composition and distribution of species, suggesting that the current species checklist is still far from being complete, not just for Lanzarote but also for the whole archipelago. In order to contribute to the knowledge of the Canary arachnofauna, we provide here the first report of the crab spider genus Heriaeus Simon, 1875 (Araneae: Thomisidae) for the Canary Islands, based on specimens of Heriaeus buffoni (Audouin, 1826) collected on this island.
Material and methods
Specimens were collected by using non-baited pitfall traps without killing-preserving agents, subsequently preserved in 99% ethanol and examined under a Zeiss Stemi 2000 stereomiscroscope. Individuals were identified to species level by examining morphological characters of the male genitalia, which is known to be a conserved character within this genus (Loerbroks 1983). In the absence of a published key of Heriaeus species, the following articles were consulted to compare male palps: Levy (1973), Loerbroks (1983), Utochkin (1985), Liang et al. (1991), Niekerk & Dippenaar-Schoeman (2013), Komnenov (2017) and Tang & Li (2010). Illustrations were made using the vector graphics editor Inkscape based on photos taken on a Canon EOS 750D camera. Microscopic images of the habitus and the pedipalp were taken using a DMC5400 camera with L.A.S. 4.13.0 software adapted to an Axio Imager M2 microscope (Carl Zeiss Microscopy) and stacked with Helicon Focus 8.1.0 (Helicon Soft Ltd). The map showing the distribution of the species was made using QGIS v. 3.16. Layers of the shape of the islands as well as the orthophotography of Arrecife were downloaded from GRAFCAN (2021). Two specimens were deposited in the collection of the Department of Animal Biology of the University of La Laguna (DZUL), another two in the invertebrate collection of the Instituto de Productos Naturales y Agrobiología (IPNA-CSIC) and one individual in the collection of the State Museum of Natural History Karlsruhe (SMNK-ARA).
Results
Heriaeus Simon, 1875
Heriaeus buffoni (Audouin, 1826) (Figs 2-3)
Specimens examined. Lanzarote, Punta Chica (Arrecife), 28.9747°N/-13.5296°W, 31. Mar. 2021, 1 ♂, M. Zarzosa leg., SMNK-ARA (19040); Punta Chica (Arrecife), 28.9771°N/-13.5301°W, 31. Mar. 2021, 1 ♂, M. Zarzosa leg., DZUL coll. (DZUL-35564); El Cable (Arrecife), 28.9577°N/-13.5752°W, 9. Apr. 2021, 1 ♂, M. Zarzosa leg., DZUL coll. (DZUL-35563); El Cable (Arrecife), 28.9543°N/-13.5765°W, 9. Apr. 2021, 1 ♂, M. Zarzosa leg., IPNA-CSIC coll.; Argana Baja (Arrecife), 28.9638°N/-13.5704°W, 16. Apr. 2021, 1 ♂, M. Zarzosa leg., IPNA-CSIC coll.
Further potential observations. Lanzarote: Femés (Yaiza), 28.9195°N/-13.7637°W, 22. Feb. 1995, 1 ex., P. Oromí obs. This specimen was not collected and thus, the identity at species-level cannot be determined with certainty.
Diagnosis. All examined specimens have a short and obtuse intermediate apophysis and their embolus tip is curved dorsally and pointed, thus fitting within the ‘buffoni-group’ after Loerbroks (1983). Heriaeus buffoni can be easily distinguished from H. buffonopsis Loerbroks, 1983 by having a larger, more massive embolus tip. Heriaeus horridus Tystshenko, 1965 (sub H. sareptanus in Loerbroks 1983) has an axe-shaped retrolateral apophysis with a step-shaped intermediate apophysis, whereas the retrolateral apophysis in H. buffoni is lanceolate with a strongly reduced intermediate apophysis.
Distribution. Algeria, Egypt, Israel (Audouin 1826, Levy 1973, Loerbroks 1983), Turkey (Kulczyński 1903) and Morocco (Nentwig et al. 2022).
Habitat. All individuals were found in nitrophilous synanthropic shrubs near urban areas. The localities of Femés, Punta Chica and Argana are composed of nitrophilous shrubs such as Caroxylon vermiculatum (L.) Akhani & Roalson, Bassia tomentosa (Lowe) Maire & Weiller and Lycium intricatum Boissier (phytocoenotic association Chenoleoideo tomentosae-Salsoletum vermiculatae) (del Arco Aguilar & Rodríguez Delgado 2018: 150). The locality of El Cable is a low scrub community settled on soils covered by a sand layer, both volcanic and organogenic, characterized by the presence of Launaea arborescens (Batt.) Murb., Cenchrus ciliaris L., Heliotropium ramosissimum (Lehm.) Sieber ex DC., Lotus lancerottensis Webb & Berthel., Polycarpaea nivea (Aiton) Webb and Pleudia aegyptiaca (L.) M.Will, N.Schmalz & Class.-Bockh. (phytocoenotic association Cenchro ciliarisLaunaeetum arborescentis) (Arco Aguilar & Rodríguez Delgado 2018: 153).
Discussion
This record of Heriaeus buffoni from Lanzarote is not only the first report of the genus for the Canary Islands, but also for any of the four Macaronesian archipelagos (Arechavaleta et al. 2005, 2010, Borges et al. 2008, 2010). The genus can be easily distinguished from any other thomisid genera currently known from Macaronesia by having the lateral eyes positioned on separate tubercles and by the densely haired body, usually covered with bristles (Levy 1985).
Heriaeus currently comprises 36 valid species inhabiting the Palaearctic and Afrotropical regions (World Spider Catalog 2021). Loerbroks (1983) classified Heriaeus species into three main groups based on their copulatory organs, i.e. the ‘hirtus-’ (Western Palaearctic including parts of North Africa), ‘transvaalicus- (South Africa) and the ‘buffoni-group’ (North Africa and Eastern Palaearctic). Within the latter group, three species are currently known: Heriaeus buffonopsis Loerbroks, 1983 (Central Asia), Heriaeus horridus Tystshenko, 1965 (Russia and Central Asia) and Heriaeus buffoni. Regarding its ecology, Lu-bin et al. (2020) collected almost 300 individuals in four different locations of the central Negev Desert Highlands (Israel), a part of the Eurasian desert belt. In Israel there are also records from the Judean Desert (Levy 1985) and in Egypt it was collected in the region of Cairo (Loerbroks 1983), but without further clarification on habitat type. Specimens in Lanzarote were also collected in desert-like ecosystems, thus fitting the known habitat on the African mainland and in Israel.
In 1995 a single individual of Heriaeus was photographed on Lanzarote (see “results”), which suggests that the species has inhabited the Canaries for at least 26 years. However, this identification was not verified with genital characters and should be treated as a tentative observation at the species level. It was first observed in Femés, in the south of the island and recently it was collected in Arrecife, in the south-eastern part of Lanzarote. Among the collected material only males were found, thus suggesting that there is a potential sampling bias and that the potentially less active females should be collected with other methods. Heriaeus buffoni does occur in Morocco (Nentwig et al. 2022), which is relatively close to Lanzarote (approximately 140 km). Given the high dispersal ability of crab spiders via ballooning (Carvalho & Cardoso 2010), it is unlikely that this species has colonized the Canaries by human assistance. Instead, a potential explanation is that it has arrived recently or that it has been overlooked in past surveys. The fact that it has been collected on two localities separated by up to 20 km, may be an indication that it is more widespread along the island. The vegetation units where H. buffoni has been collected are characteristic for most areas of the southern and central landscape of Lanzarote, thus more sampling effort in those parts of the island might uncover additional, hitherto unknown populations of the species on Lanzarote and help to determine its current distribution on the island.
Acknowledgements
The authors are indebted to Laura Brito Fernández, Cristina Calero García, Malena García Díaz, Matías Hernández González, Alejandro Leira Molina, Daniel Maldonado Egusquiza and Patricia Morera Ferrer, partners of the “Arrecife: biociudad” project of the “Arrecife Avanza” agreement, for their help during the fieldwork. In addition, we would like to thank the Canarian Government for providing a sampling permit (No Ref. Expte. 2021/17012) that allowed us to carry out this survey. We are also grateful to Tobias Bauer for his help on a preliminary version of the manuscript and for kindly taking photos of the specimen. We wish to thank Sylvain Lecigne as well as an anonymous reviewer for their constructive comments that improved the quality of the manuscript. D.S. was funded by the Ministerio de Ciencia e Innovación through an FPI PhD fellowship (PRE2018-083230) and M.Á.Z. was funded by the Arrecife local council through the program “Nuevas Oportunidades de Empleo NOE-COVID” promoted by the European Social Fund+ (ESF+).