Biodiversity Data Journal :
Taxonomic Paper
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Corresponding author: Nadhira Benhadji (nadhira.benhadji@univ-tlemcen.dz)
Academic editor: Ben Price
Received: 18 Jun 2020 | Accepted: 24 Jul 2020 | Published: 14 Aug 2020
© 2020 Nadhira Benhadji, Michel Sartori, Karima Abdellaoui Hassaine, Jean-Luc Gattolliat
This is an open access article distributed under the terms of the Creative Commons Attribution License (CC BY 4.0), which permits unrestricted use, distribution, and reproduction in any medium, provided the original author and source are credited.
Citation:
Benhadji N, Sartori M, Abdellaoui Hassaine K, Gattolliat J-L (2020) Reports of Baetidae (Ephemeroptera) species from Tafna Basin, Algeria and biogeographic affinities revealed by DNA barcoding. Biodiversity Data Journal 8: e55596. https://doi.org/10.3897/BDJ.8.e55596
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The Mediterranean basin is known to be the cradle of many endemic species. Within mayflies (Insecta, Ephemeroptera), North African species belonging to the family Baetidae remain poorly known and, traditionally, affinities to European fauna were proposed. Recent studies, based on molecular reconstructions, showed closer relationships to Mediterranean islands fauna.
Baetidae were sampled from North-West Algerian wadis (Tafna basin) and involved in COI barcoding reconstructions. Seven species were identified. The subgenus Rhodobaetis is represented by Baetis atlanticus known previously from Macaronesian islands, Europe and Morocco and the Maghrebian endemic Baetis sinespinosus. Specimens, previously identified as Cloeon cf. dipterum, correspond to Cloeon peregrinator and, until now, only reported from Macaronesia. Besides the confirmation of endemicity of some species, such as Procloen stagnicola and B. sinespinosus, our molecular study showed quite original results for relationships between European, insular and Algerian species. Baetis maurus stood out as a North African endemic sister clade to an Iberian clade. Furthermore, we found clear interspecific distances between Algerian and European clades for A. cf. sinaica and B. cf. pavidus, suggesting the presence of cryptic species in Algeria. However, additional studies are needed, as, for the moment, no clear morphological characters were found to separate the different clades and support them as valid species.
Mayflies, Baetis, Rhodobaetis, Cloeon, DNA Barcoding, COI, endemism, Algeria
The family Baetidae has a cosmopolitan distribution and represents a quarter of the Ephemeroptera diversity worldwide both at generic and specific levels (
The subgenus Rhodobaetis (corresponding to the Baetis rhodani group) presently encompasses 43 species, some of them being amongst the most common and abundant mayflies. While some species are widely distributed (e.g. Baetis rhodani (Pictet, 1843), Baetis atlanticus Soldán and Godunko, 2006), others present a presumably restricted distribution, such as endemic to a single Canary Island (Baetis palmensis Gattolliat and Sartori, 2018; B. tenerifensis Gattolliat and Sartori, 2018; B. gomerensis Gattolliat and Sartori, 2018) or known from a restricted area (Baetis chelif Soldán, Godunko and Thomas, 2005 or Baetis sinespinosus Soldán and Thomas, 1983 in Algeria) (
The species delimitation within the genus Cloeon Leach, 1815 (sensu
Recently, molecular reconstructions involving Baetidae were conducted for different projects, in particular for the origin of Macaronesian and Corsican mayflies fauna (
North African species of Labiobaetis and Cheleocloeon Wuillot and Gillies, 1993 have most probably an Afrotropical origin as they are mainly diversified in this area. A dozenspecies of Cheleocloeon are described in Afrotropics, while the genus is only represented in the Palearctic by a single Maghrebian species (Cheleocloeon dimorphicum) and one in the Arabian Peninsula, Cheleocloeon soldani Gattolliat and Sartori, 2008 (
The present study is the first molecular analysis for Algerian mayflies using the cytochrome oxidase subunit I (COI) region for species delimitation. The main aims are to clarify the status of the different species of Baetidae present in North West Algeria, especially for Central European species assumed to occur in Maghreb. We also want to clarify the species delimitation in some groups with potential cryptic species and significantdifficulties to identify, based on morphological characters only. Finally, we want to understand the affinities between Maghrebian and neighbouring fauna.
We investigated twelve sampling sites, all located in the Tafna basin in North-West Algeria (Fig.
List of COI sequenced specimens with Genbank accession number. For sites code, see
Taxa |
Sites |
Codes |
Genbank accession |
Baetis sinespinosus |
SK1 |
RB_ALG_A06 |
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SK1 |
RB_ALG_D07 |
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SK1 |
RB_ALG_E07 |
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SK1 |
RB_ALG_F05F |
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SK1 |
RB_ALG_G05F |
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SK1 |
RB_ALG_H05F |
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CH0 |
RB_ALG_D01F |
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CH1 |
RB_ALG_B03 |
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CH1 |
RB_ALG_F02 |
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CH1 |
RB_ALG_H02 |
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CH1 |
RB_ALG_A03F |
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CH1 |
RB_ALG_G02F |
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CH4 |
RB_ALG_B04 |
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CH4 |
RB_ALG_C04 |
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IOM |
RB_ALG_F04 |
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TGB |
RB_ALG_C10 |
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TGB |
RB_ALG_D10F |
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TFF |
RB_ALG_A10 |
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TFF |
RB_ALG_B10 |
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Baetis atlanticus |
SK1 |
RB_ALG_C07 |
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CH0 |
RB_ALG_C01F |
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KH1 |
RB_ALG_C08 |
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KH1 |
RB_ALG_D08 |
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KH1 |
RB_ALG_E08 |
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KH1 |
RB_ALG_H08 |
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KH1 |
RB_ALG_F08 |
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KH1 |
RB_ALG_G08F |
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Baetis cf. pavidus |
SK1 |
BP_ALG_F07 |
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SK1 |
BP_ALG_G07 |
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CH1 |
BP_ALG_B02 |
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CH1 |
BP_ALG_C02 |
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CH1 |
BP_ALG_D02 |
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CH1 |
BP_ALG_E02 |
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CH4 |
BP_ALG_A02 |
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IOM |
BP_ALG_C05 |
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TGA |
BP_ALG_D09 |
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TGA |
BP_ALG_E09 |
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TFF |
BP_ALG_F09 |
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TFF |
BP_ALG_G09 |
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TFF |
BP_ALG_H09 |
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Baetis maurus |
CH1 |
BM_ALG_G01 |
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CH1 |
BM_ALG_H10 |
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SK1 |
BM_ALG_D05F |
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SK1 |
BM_ALG_E05F |
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Acentrella cf. sinaica |
IOM |
AC-ALG-A05 |
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Cloeon peregrinator |
CH0 |
CO_ALG_B01F |
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CH1 |
CO_ALG_C03 |
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CH1 |
CO_ALG_D03 |
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SK1 |
CO_ALG_D06 |
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Procloeon stagnicola |
KH1 |
PC_ALG_A08 |
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KH1 |
PC_ALG_B08 |
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KH1 |
PC_ALG_H07 |
We performed DNA extraction using DNeasy Blood & Tissue kit (QIAGEN) and BioSprint 96 extraction robot (Qiagen) by soaking each specimen in buffer and proteinase K at 56°C for an overnight incubation. The mitochondrial DNA cytochrome oxidase c subunit I gene (COI) was amplified using the primers LCO1490 and HCO2198 (
We corrected and edited forward and reverse sequencing reads using Bioedit, then we assembled each of the two complementary sequences using Codon Code Aligner (demo mode) and obtained sequence alignments (Suppl. materials
To reconstruct the trees, we used Mega version 10.0.4; we chose the best evolutionary model using the AICc criteria (
Species delimitations
Based on the molecular analysis, seven Baetidae species were recognised or confirmed in the Tafna basin sites.
Subgenus Rhodobaetis
We obtained 11 haplogroups of Rhodobaetis (Suppl. material
Distances within (in bold) and between Rhodobaetis haplogroups. RB-Gp1: Baetis (Rhodobaetis) atlanticus; RB-Gp2: Baetis (Rhodobaetis) sinespinosus. Haplogroups with Algerian sequences are underlined.
RB_Gp1 |
RB_Gp2 |
RB_Gp3 |
RB_Gp4 |
RB_Gp5 |
RB_Gp6 |
RB_Gp7 |
RB_Gp8 |
RB_Gp9 |
RB_Gp10 |
RB_Gp11 |
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RB_Gp1 |
0.02 |
||||||||||
RB_Gp2 |
0.15 |
0.004 |
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RB_Gp3 |
0.16 |
0.14 |
n/c |
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RB_Gp4 |
0.15 |
0.14 |
0.12 |
0.004 |
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RB_Gp5 |
0.15 |
0.14 |
0.14 |
0.10 |
0.01 |
||||||
RB_Gp6 |
0.17 |
0.17 |
0.18 |
0.18 |
0.20 |
n/c |
|||||
RB_Gp7 |
0.18 |
0.16 |
0.17 |
0.14 |
0.16 |
0.12 |
n/c |
||||
RB_Gp_8 |
0.20 |
0.19 |
0.17 |
0.17 |
0.20 |
0.14 |
0.12 |
0.01 |
|||
RB_Gp_9 |
0.18 |
0.16 |
0.15 |
0.17 |
0.20 |
0.19 |
0.18 |
0.21 |
0.003 |
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RB_Gp10 |
0.19 |
0.17 |
0.17 |
0.16 |
0.17 |
0.19 |
0.18 |
0.19 |
0.10 |
0.01 |
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RB_Gp11 |
0.26 |
0.24 |
0.23 |
0.23 |
0.25 |
0.22 |
0.23 |
0.25 |
0.23 |
0.22 |
0.002 |
Baetis cf. pavidus
We obtained two haplogroups of Baetis cf. pavidus (Fig.
Distances within (in bold) and between Baetis cf. pavidus (BP_Gp1-BM_Gp2), Baetis alpinus (BP_Gp3) and Baetis lutheri haplogroups (BP_Gp4-BP_Gp7). Haplogroup with Algerian sequences is underlined.
BP_Gp1 |
BP_Gp2 |
BP_Gp3 |
BP_Gp4 |
BP_Gp5 |
BP_Gp6 |
BP_Gp7 |
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BP_Gp1 |
0.002 |
||||||
BP_Gp2 |
0.11 |
0.003 |
|||||
BP_Gp3 |
0.21 |
0.23 |
0.004 |
||||
BP_Gp4 |
0.22 |
0.24 |
0.23 |
n/c |
|||
BP_Gp5 |
0.23 |
0.25 |
0.24 |
0.08 |
0.006 |
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BP_Gp6 |
0.23 |
0.25 |
0.25 |
0.15 |
0.15 |
0.008 |
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BP_Gp7 |
0.24 |
0.24 |
0.24 |
0.23 |
0.22 |
0.23 |
n/c |
Baetis maurus
We delineated three B. maurus haplogroups (Suppl. material
Distances within (in bold) and between Baetis maurus (BM_Gp1-BM_Gp3) and Baetis cf. alpinus haplogroups (BM_Gp4-BM_Gp7). Haplogroup with Algerian sequences is underlined.
BM_Gp1 |
BM_Gp2 |
BM_Gp3 |
BM_Gp4 |
BM_Gp5 |
BM_Gp6 |
BM_Gp7 |
|
BM_Gp1 |
0 |
||||||
BM_Gp2 |
0.16 |
0.003 |
|||||
BM_Gp3 |
0.15 |
0.16 |
0.007 |
||||
BM_Gp4 |
0.24 |
0.24 |
0.23 |
0.043 |
|||
BM_Gp5 |
0.26 |
0.24 |
0.21 |
0.23 |
0.006 |
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BM_Gp6 |
0.24 |
0.23 |
0.22 |
0.25 |
0.19 |
0.06 |
|
BM_Gp7 |
0.25 |
0.26 |
0.22 |
0.25 |
0.19 |
0.19 |
0.04 |
Acentrella cf. sinaica
The reconstruction divided Acentrella sequences into 6 haplogroups (Fig.
Distances within (in bold) and between Acentrella haplogroups. AC-Gp1: Acentrella nadineae. AC-Gp2: Acentrella parvula. AC-Gp3: Acentrella turbida. AC-Gp4: Acentrella lapponica. AC-Gp5: Acentrella cf. sinaica. AC-Gp6: Acentrella sinaica. Haplogroup with Algerian sequence is underlined.
AC_Gp1 |
AC_Gp2 |
AC_Gp3 |
AC_Gp4 |
AC_Gp5 |
AC_Gp6 |
|
AC_Gp1 |
0 |
|||||
AC_Gp2 |
0.19 |
0.011 |
||||
AC_Gp3 |
0.23 |
0.21 |
0 |
|||
AC_Gp4 |
0.22 |
0.22 |
0.24 |
0.002 |
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AC_Gp5 |
0.25 |
0.25 |
0.26 |
0.22 |
n/c |
|
AC_Gp6 |
0.26 |
0.25 |
0.24 |
0.25 |
0.19 |
0.003 |
Cloeon peregrinator
We obtained 11 haplogroups from the reconstruction (Suppl. material
Distances within (in bold) and between Cloeon haplogroups. CO_Gp1-CO-Gp6: Cloeon dipterum sl. CO_Gp2: Cloeon peregrinator. Haplogroup with Algerian sequences is underlined.
CO_Gp1 |
CO_Gp2 |
CO_Gp3 |
CO_Gp4 |
CO_Gp5 |
CO_Gp6 |
CO_Gp7 |
CO_Gp8 |
CO_Gp9 |
CO_Gp10 |
CO_Gp11 |
|
CO_Gp_1 |
0.002 |
||||||||||
CO_Gp_2 |
0.09 |
0.002 |
|||||||||
CO_Gp_3 |
0.10 |
0.10 |
0.002 |
||||||||
CO_Gp_4 |
0.11 |
0.11 |
0.07 |
0.012 |
|||||||
CO_Gp_5 |
0.11 |
0.11 |
0.08 |
0.08 |
0.021 |
||||||
CO_Gp_6 |
0.09 |
0.09 |
0.1 |
0.1 |
0.1 |
0.002 |
|||||
CO_Gp_7 |
0.15 |
0.15 |
0.17 |
0.17 |
0.16 |
0.16 |
0.01 |
||||
CO_Gp_8 |
0.18 |
0.19 |
0.19 |
0.19 |
0.20 |
0.20 |
0.15 |
0.00 |
|||
CO_Gp_9 |
0.19 |
0.21 |
0.2 |
0.19 |
0.19 |
0.20 |
0.20 |
0.22 |
0.013 |
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CO_Gp_10 |
0.17 |
0.19 |
0.17 |
0.18 |
0.18 |
0.19 |
0.19 |
0.20 |
0.11 |
0.005 |
|
CO_Gp_11 |
0.17 |
0.19 |
0.16 |
0.17 |
0.17 |
0.17 |
0.18 |
0.18 |
0.18 |
0.19 |
n/c |
The CO_Gp2 haplogroup, which includes four sequences from Algeria and sequences of C. pereginator from Madeira (type locality) and Gran Canaria (Fig.
Procloeon stagnicola
In this reconstruction, six haplogroups (Fig.
Distances within (in bold) and between Procloeon haplogroups (PC_Gp1-PC_Gp5) and Centroptilum luteolum group (PC_Gp6). PC-Gp1: Procloeon stagnicola. PC-Gp2 – PC-Gp3: Procloeon bifidum. PC-Gp4: Procloeon pennulatum. Haplogroup with Algerian sequences is underlined.
PC_Gp1 |
PC_Gp2 |
PC_Gp3 |
PC_Gp4 |
PC_Gp5 |
PC_Gp6 |
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PC_Gp1 |
0.006 |
|||||
PC_Gp2 |
0.16 |
0.027 |
||||
PC_Gp3 |
0.18 |
0.19 |
n/c |
|||
PC_Gp4 |
0.19 |
0.19 |
0.20 |
0.016 |
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PC_Gp5 |
0.19 |
0.20 |
0.21 |
0.20 |
0.037 |
|
PC_Gp6 |
0.22 |
0.23 |
0.24 |
0.24 |
0.23 |
0 |
The different trees we obtained allowed us to better understand the composition of Algerian Baetidae. Based on our analysis, we can link the Algerian lineages with their sister-groups, calculate the maximum and minimum distances and evaluate which lineages may represent putative species.
Maghrebian endemic species
Baetis (Rhodobaetis) sinespinosus Soldán and Thomas, 1983
Rhodobaetis is a subgenus of Baetis and corresponds to the concept of Baetis rhodani species-group (
Procloeon stagnicola Soldán and Thomas, 1983
Our results showed a high interspecific distance between the Algerian clade and its European sister species Procloeon bifidum; thus, it confirms the validity of Procloeon stagnicola. This latter differs from P. bifidum especially by the flat and rounded bristles on the labrum margin; the pointed apex of the gills with an extremely reduced second lamella and also by the lateral margins of the abdominal segments which possess spines from segment V to IX. The species was originally described from Algeria (
Baetis maurus Kimmins, 1938
Baetis maurus is a representative of the Baetis alpinus species-group. The species is considered as an Atlanto-Mediterranean element (
Acentrella cf. sinaica Bogoescu, 1931
Acentrella sinaica is a South and Central European species (
Western Mediterranean species
Baetis cf. pavidus Grandi, 1949
Baetis pavidus is a Western Mediterranean species and was originally described from Italy and then reported from the Maghreb (
Widely-distributed species
Cloeon peregrinator Gattolliat and Sartori, 2008
Cloeon peregrinator was first considered as an endemic species from Madeira (
Baetis atlanticus Soldán and Godunko, 2006
We summarise the state of the knowledge and the implication of the present study for the Baetidae fauna of North-West Algeria in Table
Distribution of the Baetidae of the Tafna catchment. Distribution prior to study, based on 1
Operational Taxonomic Units |
Implications from this study |
Acentrella cf. sinaica |
New unnamed species in North Africa |
Baetis maurus |
Endemic to North Africa1,2; new unnamed species in Iberian Peninsula |
Baetis cf. pavidus |
North Africa and South of France1,2 |
Baetis (Rhodobaetis) atlanticus |
First report for North Africa; known from Macaronesia and Atlantic Europe1,3 |
Baetis (Rhodobaetis) sinespinosus |
Confirmation of North African endemism1 |
Cloeon peregrinator |
First report for North Africa; known from Macaronesia1 |
Procloeon stagnicola |
Confirmation of North African endemism1 |
The next steps will be to sequence more specimens from different areas of Algeria and also from Morocco and Tunisia to confirm the monophyly of the different North African clades. The results, especially the validation of the new species hypotheses, need to be confirmed by integrative methods. Only morphological evidence and more mitochondrial or nuclear genes can validate the specific status of these clades. Our study may have implications outside of North Africa, as our results suggest that one or two lineages, previously supposed to belong to Baetis maurus, may represent new species in Spain, as well as the presumably non-conspecificity of the French and Italian lineages of Baetis pavidus.
The first author would like to express her profound gratitude to Laurent Vuataz for his tremendous help and guidance with the bioinformatic treatments. The authors would like to thank also the team of the Biophore laboratory - Department of Ecology and Evolution of the Faculty of Biology and Medicine at Lausanne University: Maud Liegeois, Zoé Dumas and Karim Ghali. A special thanks to Marion Podolak of the Museum of Zoology in Lausanne for her great contributions in the biomolecular laboratory manipulations and to Laurent Vuataz for his constant help with genetic analysis.