Despite local anthropological modifications since one century, T. stagnalis, identified morphologically by Gauthier (1928) in northern Algeria still exist today. El-Frine and Réghaia are the only populations recorded to date. The genus Tanymastix has a discontinuous distribution area, it is extremely rare in Algeria, and absent from Tunisia (Marrone et al. 2016). Its occurrence is also interrupted in space in Europe (Brtek 1962; Brtek and Thiéry 1995; Brendonck et al. 2017; Thiéry 2017; Thiéry et al. 2019) and in a way, the distribution in Algeria is quite similar to that in Europe. However, in the case of Algeria, part of the discontinuity with European populations is linked to the presence of the Mediterranean Sea. This situation could suggest several scenarios on the evolution of the genus Tanymastix. Algerian populations could formerly have been separated from the European and Moroccan populations and constitute a separate species, an ancient population of T. stagnalis or a bad identified T. affinis populations which are the only other African populations of the genus (Daday 1910; Thiéry 1987; Van den Broeck et al. 2015). However, the phylogenetic analysis of the different populations of Tanymastix clearly demonstrates that populations from Algeria do not constitute an old distinct group and belongs without any context to T. stagnalis. This analysis confirms also the results of Rodríguez-Flores et al. (2019) where the Iberian Peninsula may have acted as a center of lineage accumulation with high genetic diversification between populations of T. stagnalis. According to these results, T. stagnalis species appears to have originated in the Iberian Peninsula and has colonized the rest of Europe, and more recently, the northern Algeria, while another species, T. affinis, occupied Morocco for a long time. More precisely, the Algerian populations are included in the E2 subclade, which has a large and patchy distribution through Europe, from North-Eastern Spain to Germany, with populations in France and North of Italy (Ketmaier et al. 2005; Rodriguez-Flores et al. 2019).
The strong connection between continental European and Algerian populations raises questions about dispersal methods for this genus and establishment capacity. The dispersal mechanisms of large branchiopods have been extensively documented. Studies have demonstrated that anostracans are passively scattered through their resting eggs from their habitats by various vectors such as mammals (Thiéry 1991), fishes (Beladjal et al. 2007), amphibians (Bohonak and Whiteman 1999) and insects (Beladjal and Martens 2009). Long distance colonization beyond the Mediterranean Sea implies other modes of propagation such as wind (Graham 2008), birds (Green et al. 2023) or even through human intervention (Waterkeyn et al. 2010). It should be noted that there are several bird migration roads which connect both sides of Mediterranean coasts and which can explain the colonization of Algeria by European populations rather than Moroccan (Fig. 4). However, the phylogenetical tree showed that T. stagnalis populations from Algeria are not connected with those of Corsica and Sardinia, despite a smaller geographical distance.
In the genus Branchipus, cryptic species are also distributed on both continents and some of them have a wide, interrupted distribution in Algeria and France or Italy, which is somewhat similar to the situation of T. stagnalis (Lukić et al. 2019). On the other hand, if we compare Tanymastix with Chirocephalus of diaphanus group we observe: that the distribution of the genus Chirocephalus is more continuous, that there are many clades and that there is no clade/species in common between continental Europe (France/Spain/Italy) and Algeria (Renier et al. 2013; Boumendjel et al. 2018, 2023). The phylogeographic model of Rogers (2015) would also explain this different distribution between Chirocephalus spp. and T. stagnalis by considering that Chirocephalus spp. would be a more anciently installed group in Algeria than Tanymastix and this results in a different genetic structure with more localized lineages. Thus, the “Monopolization Hypothesis” would explain the strong priority effects of the founding population, which, after installation, would hinder the gene flow of newcomers occupying the same niche (Meester et al. 2002). In this case, the success of the European Tanymastix would be linked to the absence of indigenous Tanymatix species in the north of Algeria, and therefore, without local competition, they could easily establish themselves, while, in the case of Chirocephalus, the European populations/species are faced with very strong local competition and do not take root (Boumendjel et al. 2018, 2023). Furthermore, the presence of a consistent cystbank of the resident species, which are adapted to local conditions, limits the establishment of the immigrant species (Rogers et al. 2015).
Ecological notes
Algerian T. stagnalis populations live in arable sandy pools located in shady areas, with subhumid/humid climate close to the Mediterranean Coast. This type of habitats, especially ponds of costal sand bar, is very similar to that of Spanish populations (Olmo et al. 2015). T. stagnalis has been reported to live in temporary pools on sandy soil (Brtek and Thiéry 1995; Zarattini et al. 2017), karst systems (Mura and Zarattini 2000), rock pools (Rabet 1994; 2001), pools in forests (Gauthier 1928; Rabet 2001) and also in tree holes (Jocque et al. 2013). Ponds similar to those of El-Frine are numerous in the humid and sub-humid regions of the country but they harbour Chirocephalus salinus or C. sanhadjaensis (Samraoui and Dumont 2002; Boumendjel et al. 2018) rather than T. stagnalis.
The species develops in Algeria in winter until early spring. It was recorded from December to April at temperatures between 14°C and 22°C in Réghaia pools (Gauthier 1928). In Spain it develops in spring only (Olmo et al. 2015), while in Italy, it was found in spring after melting snow (Mura and Zarattini 2000) and also, in autumn after the first rains (Zarattini et al. 2017). In France, it occurs throughout the year in the rock pool of the Fontainbleau forest (Rabet 2001). In fact, T. stagnalis was first classified as a cold stenotherm species (Fløssner 1972, Grainger 1991), but Eder et al. (1997) suggested that the species should be considered a warm stenotherm, since it has been found at temperatures above 30°C in Austria. The large temperature interval of its occurrence between North-African and European climate suggests rather a eurytherm species. T. stagnalis prefers freshwater with very low conductivity. It was recorded with a maximum of 0.8 mS.cm− 1 in El-Frine, which agrees with the results of several authors (Alonso 1985; Brtek and Thiéry 1995; Rabet 2001; Wood 2002; Cancela da Fonseca et al. 2008; Arukwe and Langeland 2013). Occasionally, a threshold of 7.29 mS.cm− 1 has been registered for adults by Rueda-Sevilla et al. (2006), at the end of the hydroperiod. The pH in El-Frine was neutral to alkaline. Gauthier (1928), Rabet (2001) and Arukwe and Langeland (2013) reported pH around neutrality (6.5–7.2), probably because of punctual studies. The survey of Olmo et al. (2015) showed neutral to alkaline pH (up to 9), while that of Zarattini et al. (2017) resulted in acidic to alkaline pH, with a threshold of 4.9. Dissolved oxygen values are extreme, showing the wide range adaptation that can occur in small temporary pools (Brtek and Thiéry 1995; Boix 2002; Olmo et al. 2015).
Individual density of T. stagnalis of El Frine was close to that of Spanish or Italian populations. Olmo et al. (2015) reported densities comprised between 0.1 and 11 ind.L− 1for Spain, while Zarattini et al. (2017) reported higher densities between 2 and 49 ind.L− 1 for Italy. Vanschoenwinkel et al. (2013) considered a density of 2 ind.L− 1 of Branchipus schaefferi from ponds on arable lands as high, which sounds a good establishment of the species, despite the agricultural pressure, including pesticide and fertilisation.
All developing stages were often present in El-Frine and males were dominant, as observed in other anostracans living in harsh conditions (Beladjal et al. 2003; Amarouayache et al. 2010). This male biased sex-ratio is encountered in several anostracan species; it favours the genes mixing to limit consanguinity in new generations and contribute to species persistence (Lievens et al. 2016). Egg bank is an important criterion to take into account in anostracan assessment (Brendonck and De Meester 2003). T. stagnalis resting eggs number of 0.35 eggs.cm− 3 (35230 eggs.m− 2) found in El-Frine is relatively weak, and comparable to that of the “inactive” Romanian population (Demeter and Peter 2011). It was lower than that of the Italian population (5.7 eggs.cm− 3) and much lower than that of B. schaefferi (761.9 eggs.cm− 3) (Zarattini et al. 2017). Thiéry (1997) reported more than 170000 eggs.m− 2 for C. diaphanus in ditches at Rocheford du Gard in France.
The co-habitation of T. stagnalis with Atyaephyra desmaresti is reported for the first time. T. stagnalis was found to co-habit with two other large branchiopods, in several ponds of El-Frine region, Chirocephalus salinus (dishes) and rarely with Cyzicus tetracerus (Krynicki 1830) (Ghaouaci 2018). In Réghaia, T. stagnalis co-habits with C. algeriensis and L. apus lubboki, which appeared later in the rearing medium. In Southern Europe, T. stagnalis co-habits with two anostracans C. diaphanus and B. schaefferi in Southern France (Camargue) (Nourisson and Aguesse 1961) and in Italy (Mura 1991). Waterkeyn et al. (2011) showed that the Notostracan Triops cancriformis predates on T. stagnalis cysts. We notice that L. apus is also an active predator of T. stagnalis adults (unpublished data). The other associated crustacean fauna (copepods, cladocerans and ostracods) is quite similar in both regions, Réghaia and El-Frine (Gauthier 1928; Samraoui and Dumont 2002; Ghaouaci et al. 2017, 2018).
Ecological preferences of T. stagnalis are not clearly understood, and despite its broad environmental tolerance limits, the species do not appear to establish in close biotopes with similar physico-chemical and edaphic conditions (Grainer 1991). Other biological factors should curb establishment of the species in new habitat, by inhibiting egg hatching, among them, chemical compounds produced by resident species, such as faeces, micro-algae (Griffiths et al. 1991) or pheromones (Beladjal et al. 2007).
El-Frine population is stable, develops throughout the entire hydroperiod and several ponds confined there between Oubeira and Tonga Lakes harbour the species on a perimeter of 1.6 km², at least since works of Gauthier (1928). Considering the limited sites in Algeria and the risks of its disappearance due to global warming and anthropological pressure, the species should be preserved and assessed among the endangered species on the IUCN red-list.