Tropical forests that cover less than 7% of the planet are disappearing day by day. In Africa, the degradation of forest cover and the loss of biodiversity has become very worrying with alarming proportions. Africa's forest cover has decreased from 749 million hectares in 1990 to 674 million hectares in 2010 [1] In Côte d'Ivoire, a country located in West Africa, forest cover is also decreasing drastically under the influence of climate change and anthropogenic actions. Forest cover has decreased from over 16 million hectares in the 1900s to 1,385 million hectares in 2010 [2] [3]Along with the forests, a great diversity of animal and plant life forms, divided into four ecological zones of the country, is also disappearing with the forests [4-5]The dissemination of knowledge on spiders in Côte d'Ivoire has been done by a dozen authors. The publications of some of them are based on the ecology of spiders while those of others consist in describing the specimens. A total of 20 publications have been listed. Works on spiders in Côte d'Ivoire were carried out in the center; east, west and north of the country (see Fig. I). This article is a literature review which, on the one hand, gives the history and location of the collection areas of spiders in Côte d'Ivoire, and on the other hand, presents the authors and the spiders collected and described. This synthesis of previous works will help and enable successful of future studies within the country. It is a checklist of known species recorded in the country. 

The studies about spiders in Côte d'Ivoire began in 1961 with research works done at Lamto by Maxime Lamotte, Yves Gillon and Dominique Gillon [6] The monitoring of spiders in Lamto savannah occured from 1962 to 1965. It was performed by a team led by Yves and Dominique Gillon. Overall, more than 100,000 spiders were collected during these samplings [6] Over this same period from 1962 to 1965, the researcher Jean-François Jézéquel had participated in the collection of spiders from Lamto savannah between 1962 and 1963. He had also collected spiders in Yealé, a village at Mount Nimba’s foothill. Mr. Paul Planquette also took part in collecting spiders from Lamto savannah between 1963 and 1964 [7-8] In 1969, Maxime Lamotte proposed a thesis topic on the ecology of spiders in Lamto savannah. After Patrick Blandin's first results, Marie-Louise Célérier, a specialist in spider energy balances, was associated to establish energy balances for a few representative species of Lamto savannah [6] Later in 2005, during a mission to assess the quality of forest fragments’ biodiversity in the eastern region of Côte d'Ivoire, Rudy Jocque collected spiders [9] To date, the only Ivorian arachnologist is Professor Bessekon Denis ASSI. He worked in 1992 at the University of Nancy 1 in France, on the contribution to the study of the mechanisms involved in the regulation of maternal behaviour in the Coelotes terrestris spider (Araneae, Agelenidae) [10] Table I provides a summary of the timelines and locations where the researchers worked in the country. 

Geographical distribution of spider collection areas in Côte d'Ivoire 

In Côte d'Ivoire, a total of 331 species of spiders belonging to 168 genera and 46 families have been identified. The families were mostly collected in [11-13] the central part of the country, in Lamto pre-forest savannah, between Tiassalé and Toumodi; in the listed forests in the East of the country: Bossematié (Abengourou, Appouesso), Mabi (Bettie, Mebifon) and Songan (Bettie); in the Center, at Bouaké; in the North, in Korhogo and in the West, at Mount Nimba in Tai National Park.

Table I. Geographical coordinates of spider collection points in Côte d'Ivoire

Figure Image is available at PDF file

Figure I. Geographical distribution of spider collection areas in Côte d'Ivoire

Various works carried out in Côte d'Ivoire

The first document on spiders in Côte d’Ivoire has been written in 1964 by Jean-François Jézéquel on description of the new spider species from Lamto savannah. The second documents were published by Yves Gillon and Dominique Gillon in 1967.  Just after these authors, in 1971, Patrick Blandin began writing about the same subject [14,5]. 

However, from a biogeographical point of view, the ecological studies taking into account data from savannahs and forest galleries, which have enabled a complete inventory and characterization of Lamto's pre-forest savannah spiders in Côte d'Ivoire, began with works of Blandin and Célérier in 1981. But it is important to note that this study could be carried out by the research initiative of Maxime Lamotte, Yves and Dominique Gillon since 1961 [5,14]

The studies carried out so far on spiders in Côte d'Ivoire are summarized in chronological order. The main part of these writings contains the site and the period of sampling, the species and families notified in the basic documents. 

First of all, the methodologies used by the various authors are as follows:

• Threshing and excavation of trees and shrubs [7,8,15]

• Quadra trapping, pit trap and mowing [16-20]

• Distance sampling based on the reflective characteristics of the eyes [21,22]

[7] this study was carried out at Lamto in Singrobo savannah between Tiassalé and Toumodi, from 1962 to 1963. During this period, he captured several tens of thousands of spiders. Only hundreds of the most abundant spider species were analyzed. Rare species have been described. The descriptions in this document begin with one of the species (Scytodes reticulata) of the family Sicariidae.   

[23] in this document, the author studied some new species collected at Lamto, near Singrobo, from 1962 to 963. He described species from two families (Palpimanidae and Zodariidae). These families were represented by few species, but abundant in numbers of individuals encountered in various herbaceous environments, shrubs and roosts. Thus, it described eight species belonging to four different genera. For the family Palpimanidae, he obtained five species: genus Scelidocteus, Simon 1907 (Scelidocteus lamottei and Scelidocteus vuattouxi) and genus Sarascelis, Simon 1887 (Sarascelis lamtoensis, Sarascelis rebiereae and Sarascelis junquai). The author then described the species of the family Zodariidae: genus Storena, Walckenaer 1805 (Storena bandamaensis and Storena bicolor) and genus Chariobas, Simon 1893 (Chariobas cylindraceus).

[15] this study was a continuation of the previous systematics. He described eight new species belonging to the family Drassidae. These spiders were also collected in Lamto savannah, particulary in the trees and shrubs. The author described the following species: genus Drassodes, Westring 1851 (Drassodes falciger), genus Odontodrassus, Jézéquel 1965 (Odontodrassus nigritibialis and Odontodrassus bicolor), genus Echemus, Simon 1878 (Echemus incinclus), genus Zelotes, Gistel 1848 (Zelotes singroboensis), genus Aphantaulax, Simon 1878 (Aphantaulax cincta), genus Scotophaeus, Simon 1893 (Scotophaeus crinitus), genus Nomisia, Dalmas 1921 (Nomisia uncinata and Nomisia eburneensis). He also reported the presence of an Abyssinian species of the genus Xerophaeus, Purcell 1907 (Xerophaeus coruscus) that had never been recorded in West Africa. 

[8] this systematic study was a continuation of his previous work. The description of the new species was about the family Thomisidae. The majority of individuals of this family were collected at Lamto, from 1963 to 1964 by Mr. Paul Planquette. Some individuals were sampled in the gallery forests of Lamto and other individuals collected in Yealé, at the foothill of Mount Nimba, by Jézéquel himself.  He described ten species which are: genus Stiphropus, Gerstaeeker 1873 (Stiphropus melas), genus Dieta, Simon 1880 (Dieta ribes), genus Tmarus, Simon 1875 (Tmarus planquettei), genus Pherecydes, Pick. Cambr.O. 1883 (Pherecydes zebra), genus Xysticus, Koch 1835 (Xysticus argenteus and Synaema viridisternis), genus Firmicus, Simon 1895 (Firmicus aurantipes), genus Gephyra, Koch 1874 (Gephyra glauca), genus Thomisops, Jézéquel 1964 (Thomisops cretaceus), genus Parasmodix, Jézéquel 1966 (Parasmodix quadrituberculatus). 

[17] these authors conducted an ecological study to determine the annual cycles of arthropod numbers and biomasses in the herbaceous stratum of the Lamto savannah, from 1964 to 1966. Spiders were sampled during the long dry season, the short dry season and the long rainy season. Biomasses and numbers were also jointly evaluated according to the period of fire.

Among these arthropods, they mention that the biomass and numbers of spiders varied positively during the year. Spiders adapted very quickly to the changes of seasons and the action of fire.

[16] this study consisted in comparing invertebrate population of Lamto savannah in Côte d'Ivoire to those of Fété-Olé in the Sahel in northern Senegal. The objective of this work, carried out at Lamto in 1965 and at Fété-Olé in 1971, was to assess the limits of variation of savannah fauna of invertebrates. Similarly, LAMTO and Fété-Olé present the spider population as the most abundant group of arthropods. The spiders, more abundant in Lamto than in Fété-Olé, were therefore the most abundant carnivores of the two environments. In the Lamto savannah, the spiders were more abundant after the passage of the fire than before, because the organisms’ inhabitants of the vegetal cover finding themselves on the ground without support and without shelter, were prey for the spiders which would then leave their refuges in the ground in greater numbers than usual.

[18] This document dealed with a preliminary general observation on Lamto spiders, regularly collected just after the fire of December 30, 1961 until 1965 by a team led by Yves and Dominique Gillon. The sampling took place in the burnt savannah (BS) and unburnt savannah (UBS). Spiders from these environments were collected during the long dry season (LDS), the long rainy season (LRS), the short dry season (SDS) and the short rainy season (SRS). It appeared that the Lycosidae family (about 25 to 34%) was the most dominant, regardless of the type of environment. This abundance was followed by Ctenidae (7-15%), Salticidae (11%), Thomisidae (8-9%) and Clubionidae plus Eusparassidae (6-9%).

In total, the representatives of five families (Lycosidae, Salticidae, Thomisidae, Clubionidae and Eusparassidae (including Cteninae)) thus constitute 60 to 70 % of the spider population and the remaining 30 to 40% of the spiders correspond to the representatives of about 15 families. Depending on the season, Lycosidae always had a dominant frequency and remained constant regardless of the season. The Salticidae stood out during the two short dry and rainy seasons. They represented nearly 20 % of the population during the short dry season while the frequencies of several families such as Thomisidae, Sicariidae, Pisauridae and Drassidae decreased considerably. During the short rainy season, the Salticidae made up barely 10 % of the population, while practically all the other families became more and more abundant. 

However, the density of spiders collected in burnt savannah remained low compare to that of savannah protected from fires.

[19] the author depicts again with more precision the Runcinia (Runciniopsis) erythrina species. This species belongs to the family Thomisidae. It was described in 1964 by Jézéquel from a single specimen. This time, five specimens were observed with the presence of a male and an adult female. 

[20] Patrick Blandin worked by estimating the numbers and biomasses of spiders through the realization of sampling techniques (trapping by quadrats and mowing). As for Marie-Louise Célérier, she worked by applying breeding techniques to appreciate certain characteristics of some spiders, while establishing their energy balance.

Sampling carried out at Lamto for this purpose took place in four different types of savannahs: 

• savannahs burned during the previous great dry season (BS);

• savannahs protected from the fires of the previous great dry season (UBS); 

• savannas protected from fires at least during the two previous great dry seasons (SNNB);

• savannahs protected from the fires of January 18, 1965 and burned late on April 7, 1965 (SBT). 

A total of 34 spider families have been identified. The family which was more represented with more abundant specimens (20.5%), was Lycosidae. This family is followed by Salticidae (17.4%) and Thomisidae (14.5%). The other 31 families had frequencies below 10%. Four new species (Vuattouxia kouassikonani (Pisauridae); Hippasa lamtoensis, Orinocosa celerierae, Brevilabus gillonorum (Lycosidae)) were described by Blandin and his collaborators.

The determination of the individual energy balance of six species allowed assessing the trophic flow through spider populations. These species, divided into three different families, were confined in the table below:

Table II. Different species bred by M. L. Célérier.

[14] the methods of quantitative surveys, trapping, mowing and sight hunting have allowed to collect more than 200,000 spiders. These are divided into 34 families. Blandin and his collaborators have described four (4) new species belonging to two of these 34 families. These species were divided between the families Pisauridae and Lycosidae. The species Vuattouxia kouassikonani (Pisauridae) was described by Blandin himself and the other species (Hippasa lamtoensis, Orinocosa celerierae, Brevilabus gillonorum) of the family Lycosidae, were described by his collaborators.

The remarkable presence of the Lycosidae family in the Lamto savannah called the attention of the authors. They chosed the species Brevilabus gillonorum Comic, 1980, the most abundant one of the Lycosidae family for breeding. The species Brevilabus gillonorum, Comic, 1980 is univoltine, i.e. it reproduces only once a year. 

[24] the study of demographic strategies in spiders was reflected in the reproductive effort (ER'i), demographic profiles and mortality factors during rearing. The authors worked with breeding data of six species of the family Lycosidae. Four species of Lamto [20] Hippasa lamtoensis, Pardosa injucunda, Orinocosa celerierae, Brevilabus gillonorum and two other species: Pardosa lugubris, Geolycosa godeffroyi belonging respectively to the data of Edgar (1971 a, b, c) and Humphreys (1976).

The authors pointed out that in spiders, two species of the same family may have different adaptation strategies (k and r). Two species of the Lycosidae family, Revilabus gillonorum and Geolycosa godeffroyi belonged to strategists k and r, respectively. 

They described the difficulties of this study. One of the difficulties was that it is not easy to select individuals from the same cohort alone. The other difficulty was related to the environmental factors that induce mortality in spider breeding. 

[6], this article on Spiders of the Lamto savannah, is a summary publication. The author discussed the new species as Vuattouxia kouassikonani (Pisauridae) of spiders described in Lamto, mentioned in Blandin, 1983.

[25] the author described a new species: Hippasa lamtoensis Dresco (Lycosidae). This new species belongs to the genus Hippasa described in 1885 by Simon. It came from Lamto and provided by M. L. Célérier who bred it for energy balance work. The male and the female are well distinguished. 

[22] they made a revision of the genus Hippasa (Lycosidae, Araneae), an African spider. The recognition characteristics of some species of Lycosidae of the genus Hippasa were described. These species are : Hippasa affinis Lessert, 1933; Hippasa funerea Lessert, 1925; Hippasa albopunctata Thorell, 1899; Hippasa australis Lawrence, 1927; Hippasa cinerea Simon, 1898a;  Hippasa lamtoensis Dresco, 1981; Hippasa marginata Roewer, 1960; Hippasa decemnotata Simon, 1910; Hippasa natalica Roewer, 1960; Hippasa innesi Simon, 1889a; Hippasa africana Roewer, 1960.

However, three new species of the same genus have been described. This is the case of Hippasa brechti, a new species from the Lamto savannah; Hippasa elienae, a new species from the forest reserve of Tanzania and Kenya; Hippasa sinai, a new species from Egypt, Sinai and Saudi Arabia. 

[9] the evaluation of the conservation status or the assessment of the quality of the forest ecosystems of eastern Côte d'Ivoire was made possible by the nocturnal capture of more than 2600 spiders of the family Ctenidae. It appears that the family Ctenidae captured in degraded forest fragments were only half of those captured in less anthropized forest fragments. This family of spider appears to be a good indicator of ecosystem health.

[9] Lycosidae are a family of spiders, typical of degraded, open or grass-filled environments (Poaceae). They are absent where the forest canopy is closed, particulary in less anthropized forest habitats. They are also found in meadows and swampy places. They have three claws unlike the Ctenidae which have tufts of claws. The claws allow them to move and hunt in their different living environments. The presence of the two families (Lycosidae and Ctenidae) in a set of landscape could be an excellent means of assessing the quality of habitats. Thus, the proportion of Lycosidae / Ctenidae could be an excellent and easy-to-use indicator for assessing forest quality. 

[21] A total of 2607 specimens of Ctenidae were collected. Among these specimens, 15 species have been described and seven (7) of them were new. A key for the determination of these species was developed subsequently. The seven new species were: Africactenus decorosus des Arts (Arts, 1912); Africactenus evadens n. sp. [21]; Africactenus monitor n. sp. [21] Ctenus fallax n. sp.; Ctenus undulatus n. sp. [21]Ctenusperguianus des Arts (Benoit, 1980); Ctenus sp. They were followed by the 13 others: Petaloctenus bossema ; Petaloctenus songan ; Anahita aculeata (Simon, 1897); AnahitaIineata (Simon, 1897); Ctenus eminens des Arts (Arts, 1912); Ctenus kingsleyi F.O.P.- Cambridge; Ctenus mirificus des Arts (Arts, 1912); Cramsoccidentalis F.O.P.-Cambridge.

[11] they described three new species of the genus Acanthinozodium (Denis, 1966), of the family Zodariidae. These three species and their origins were: Acanthinozodium crateriferum (Ethiopia), Acanthinozodium sahelense (Sahel region) and Acanthinozodium quercicola (Morocco). In Côte d'Ivoire, three specimens of Acanthinozodium sahelense were caught by pit traps: a male and a female in riparian forests along the Bandama River in Korhogo and a male in a rice field in Bouaké. 

[12] Work on spiders on Mount Nimba contributed to the sampling of the family Ctenidae. Molecular and morphological analysis of 41 individuals of Ctenidae, including 33 from Mount Nimba, led to the setting of five new genera (Amicactenus, Macroctenus, Nimbanahita, Perictenus and Piloctenus). These analyses also revealed that 22 species were typical of Mount Nimba. Among the identified species, eight new ones have been described: Amicactenus mysticus; Macroctenus herbicola; Macroctenus nimba; Macroctenus vandenspiegeli; Nimbanahita montivaga; Perictenus molecula; Piloctenus haematostoma; Piloctenus gryseelsi. The majority of the described species come from the part of Mount Nimba originating from Guinea, only the species Piloctenus haematostoma has been collected in Tai, Côte d'Ivoire.

Families of the spiders in Côte d’Ivoire

In Côte d'Ivoire, 331 species, 168 genera and 46 families have been identified. These families are represented in the Centre (Lamto reserve) and East (classified forest of Bossematié, Mabi and Songan) [26,13,20]

A summary table of spider families in Côte d'Ivoire is established (Table III). This may have been possible thanks to the work of Blandin and Celerier (1981) and a list provided by Dippenaar-Schoeman and Jocque in 2018. 

A total of 46 families were registered. [20] provided 34 families of which eight families (Agelenidae, Clubionidae, Ctenizidae, Dictynidae, Dysderidae, Mimetidae, Oecobiidae, and Telemidae) are specific to their work. For [26] 38 families were provided including 12 families (Cithaeronidae, Corinnidae, Cyrtaucheniidae, Dipluridae, Eutichuridae, Hahniidae, Liocranidae, Philodromidae, Sicariidae, Symphytognathidae, Trachelidae, Zorocratidae) specific to these data.

Table III. List of spider families in Côte d'Ivoire (Blandin and Celerier, 1981; Dippenaar-Schoeman and Jocque, 2018). X: present and O: absent

All the works on spiders of Côte d'Ivoire, carried out by the several authors, show difficulties related to the determination or identification of species due to the lack of identification keys or appropriate documentation. The whole works revealed a total of 331 species of spiders, with 168 genera and 46 families. Among these specimens, 44 new species were described in Côte d'Ivoire over the last 20 decades. Jean-François Jézéquel, Patrick Blandin and his collaborators, Dresco described respectively 28, four and one species. Rudy Jocqué and collaborators described 11 new species. Patrick Blandin and Rudy Jocqué revised some genera and species previously described by Jézéquel. They also described other genera and species from elsewhere. In view of the anthropic pressure and climate change that contribute to the modification of natural environments, several studies on spiders are necessary in order to identify and characterize spiders likely to be of paramount importance for biological control.

Acknowledgement

This article summarizing the works carried out on spiders in Côte d’Ivoire is the result of a bibliographical research. We thank Professors Konaté Souleymane (University NANGUI ABROGOUA) and Denis Assi Bessekon (University Félix Houphouet Boigny) for their documentary contributions. Our thanks also to Professor Rudy Jocqué (Department of Zoology, Royal Museum for Central Africa, Belgium), Professor Ansie Dippenaar-Schoeman (ARC-Plant Health and Protection, South Africa) for their perfect collaboration and all the orientations and basic documentation that facilitated the completion of this work. Many thanks to Doctor Zon Déméango Serge, for his advice and guidance.

The authors declare that they have no conflict of interest

No funding sources

The study was approved by the University Nangui Abrogoua

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