Introduction

Owing to diverse topography, geology, different climatic zones and most importantly the presence of high mountains, Morocco has the most diverse butterfly fauna in the Maghreb region with a total of 136 species recorded (combined from TENNENT, 1996; TARRIER & DELACRE, 2008). The bulk of the diversity is limited to the high mountains, particularly in the Rif and Middle Atlas in the north, which receive more precipitation and have many faunistic similarities with the nearby Iberian Peninsula. Further south, the fauna of the Anti Atlas is somewhat impoverished, but still includes several Mediterranean elements, especially at higher altitudes (TARRIER, 1996; TARRIER, 1997; TARRIER & DELACRE, 2008).

Based on exhaustive studies of Michel Tarrier one could conclude that the fauna of butterflies of Morocco, and the Anti Atlas in particular, is well studied apart from the skippers which were entirely neglected in his voluminous work. However, little precise faunistic information has been published on the butterflies of the Anti Atlas so far (DE PRINS et al., 1984; BOZANO & GIACOMAZZO, 1988; MÉRIT, 2014) or only vague descriptions of the localities are provided (e.g. TARRIER, 1995, 2000, 2011). In order to fill this gap we give detailed information on the distribution of the butterflies, including skippers, in the Anti-Atlas region gathered during recent surveys. The more important finds, including one new record for the country, are discussed in detail.

Material and methods

In the last 10 years eight independent visits by the three authors were made to the southern Anti- Atlas and its surroundings in order to study the butterfly fauna. The region was first visited by MR as early as 2001. The selection of sites and areas with potentially important butterfly habitat for the more recent studies was made before the trips with the aid of Google Earth images. Adult butterflies were either netted using entomological nets and released after identification, or identified in nature without capture.

List of localitiesFig. 3

1. 1. 1

   . Taroudant, Tioute, within the village oasis, 440 m, 30º 23’ 39.21”N, 8º 41’ 57.72”W, oasis with mosaic of small fields and bushy hedges, 3-IV-2013

2. 2. 2

   . Ouzzoun, Tiferki, along the road from Taliouline to Igherm, 1300 m, 30º 13’ 30”N, 8º 16’ 29”W, dry, bushy slopes, small gullies, 14-IV-2011

3. 3. 3

   . Igherm, high plateau on the road 7037 about 2 km east of Adar village, 1810 m, 30º 7’ 41.82”N, 8º 21’ 4.00”W, rocky pasture, 10-IV-2009, 14-IV-2011

4. 4. 4

   . Tata, Taangoumte, road verge in oasis at the SE edge of the village, 1200 m, 30º 1’ 12.73”N, 8º 18’ 22.52”W, rocky road side, sandy meadows, 18-III-2015

5. 5. 5

   . Igherm, small valley on the road 7037 about 4 km northeast of Igherm, 1840 m, 30º 6’ 29.10”N, 8º 26’ 6.84”W, woody gully with dry grasslands, 10-IV-2009

6. 6. 6

   . Igherm, along the road 7038 about 6 km west of the town, 1750 m, 30º 4’ 35.76”N, 8º 31’ 9.96”W, dry grasslands, abandoned arable land, orchards, 10-IV-2009

7. 7. 7

   . Tafraoute, Amzawr, in a small valley NE of the village, 1580 m, 29º 49’ 53.48”N, 8º 44’ 32.26”W, rocky meadows, flowery banks of a dry river, 14-IV-2011, 17-III-2015

8. 8. 8

   . Tafraoute, Imi n’ Guerdane, side valley at the village, north of Ait-Abdallah, 1500 m, 29º 50’ 11.94”N, 8º 45’ 59.14”W, rocky slopes, small meadows, 10-IV-2009, 17-III-2015

9. 9. 9

   . Tafraoute, Azgour, dry riverbed and slopes W of the main road SW of the village, 1530 m, 29º 46’ 13.07”N, 8º 47’ 16.79”W, rocky meadows and slopes, dry riverbed, 15-III-2001, 16-III- 2001, 1-IV-2013, 6-IV-2013, 17-III-2015, 25-III-2017

10. 10. 10

    . Tafraoute, Touli, small lateral valleys N of the main road above the village, 1620 m, 29º 45’ 17.81”N, 8º 47’ 17.05”W, rocky meadows and slopes, dry riverbed, 25-III-2017

11. 11. 11

    . Tafraoute, Titke, small valley N of the village, 1530 m, 29º 45’ 2.07”N, 8º 48’ 54.17”W, rocky slopes, dry gully, 1-IV-2013, 6-IV-2013

12. 12. 12

    . Tafraoute, Tizi’ N Tarakatin, in the valley NW of the village, 1360 m, 29º 46’ 56.95”N, 8º 51’ 59.34”W, dry river bed, 14-IV-2011, 6-IV-2013, 25-III-2017

13. 13. 13

    . Tafraoute, SW side of the pass on the route 7056 before Tanalt, 1710 m, 29º 48’ 12.65”N, 9º 5’ 2.83”W, shrubby meadows with dwarf bushes, 8-IV-2009, 17-III-2015

14. 14. 14

    . Tafraoute, Ait Yiftan, in a small valley at the village, 1490 m, 29º 48’ 54.35”N, 9º 2’ 30.42”W, grassy meadows and orchards, 6-IV-2013

15. 15. 15

    . Tafraoute, north of Djebl Lekst on the road 7056 around the waterfall visible below the road, 1380 m, 29º 50’ 6.58”N, 9º 1’ 28.65”W, cultivated meadows, bushes, roadside, 8-IV-2009

16. 16. 16

    . Tafraoute, Ida Ougnidif, S facing slopes above the road SW of the town, 1380 m, 29º 50’ 29.86”N, 9º 1’ 30.20”W, bushy and rocky slopes, 6-IV-2013, 17-III-2015

17. 17. 17

    .Tafraoute, narrow valley few km before Ait-Iftene on the route 7056 from Ida-ou-Gnidif, 1290 m, 29º 50’ 42.36”N, 9º 0’ 40.26”W, abandoned meadows, rocky slopes, 8-IV-2009

18. 18. 18

    . Tafraoute, Tiguissas, along sideroad at the bridge N of the village, 1160 m, 29º 52’ 28.17”N, 9º 0’ 36.60”W, ruderal area, road verge, 17-III-2015

19. 19. 19

    . Tafraoute, Imhiln, slopes above main road NW of the village, 1300 m, 29º 55’ 35.40”N, 9º 0’ 32.52”W, dry grassy rocky slopes, 15-III-2001

20. 20. 20

    . Tafraoute, small hill almost encircled by the road on the route 105 some 30 km SE of Ait- Baha, 1200 m, 29º 58’ 9.15”N, 9º 1’ 10.82”W, dry grassland on sandy ground, 8-IV-2009

21. 21. 21

    . Ait Baha, Azour’n’Ali, at the turn for the village, 960 m, 30º 1’ 36.61”N, 9º 2’ 40.14”W, grassy meadows in dry hills, 1-IV-2013

22. 22. 22

    . Ait Baha, Tlata Uoanass, in the small side valley near the reservoir, 620 m, 30º 3’ 45.41”N, 9º6’ 51.89”W, grasses in dry hills, 15-III-2001

23. 23. 23

    . Ait Baha, Targua N’Touchka, oasis and the E side valley, 450 m, 29º 52’ 39.52”N, 9º 11’ 26.08”W, arable land, bushes, gravels, 26-II-2012

24. 24. 24

    . Ait Baha, Ait Bibi, the pastures along the river NE of the village, 190 m, 29º 51’ 47.03”N, 9º 17’ 20.08”W, pastures, dry grasslands, arable land, 26-II-2012

25. 25. 25

    . Tafraoute, small valley 5 km south of Agard Oudad, at the beginning of the road to Ait- Mansour, 1190 m, 29º 39’ 32.31”N, 8º 57’ 26.01”W, dry river bed, rocky slopes, 9-IV-2009

26. 26. 26

    . Tafraoute, north side of the pass north of Tlatat Tasrirt town, 1540 m, 29º 37’ 56.88”N, 8º 56’ 31.95”W, pastures, rocky slopes, 9-IV-2009, 23-III-2017

27. 27. 27

    . Tafraoute, Taghaout, in the gorge SE of the village, 1430 m, 29º 36’ 21.19”N, 8º 50’ 51.77”W, small gorge, rocky slopes and meadows, 16-III-2015, 23-III-2017

28. 28. 28

    .Tafraoute, first part of the gorge at the settlement Ait-Mansour, 1260 m, 29º 32’ 54.58”N, 8º 52’ 45.26”W, dry rocky slopes, meadows in the palmery, 9-IV-2009

29. 29. 29

    . Tafraoute, Ida Ouaassam, along small track on N facing slopes of the mountain N of the village, 1450 m, 29º 31’ 49.00”N, 9º 6’ 6.30”W, rocky and bushy slopes, 16-III-2015

30. 30. 30

    . Tafraoute, Col du Kerdous, on the ridge S of the hotel, 1250 m, 29º 32’ 45.72”N, 9º 20’ 21.04”W, rocky slopes with sparse vegetation, 16-III-2015

31. 31. 31

    . Tafraoute, Col du Kerdous, along the road down from the pass on N slopes, 1050 m, 29º 32’ 50.51”N, 9º 21’ 25.88”W, pine plantation, rocky slopes, terraces with cultivations, 16-III-2015

32. 32. 32

    . Tiznit, Sidi Ahmed Ou Moussa, dry river NE of the town, 450 m, 29º 32’ 23.38”N, 9º 26’ 49.09”W, olive trees plantation with artificial irrigation, 20-III-2016

33. 33. 33

    . Tiznit, Mirght, pastures north of the village, 830 m, 29º 24’ 17.54”N, 9º 43’ 18.81”W, small wadi with sparse almond trees, 20-III-2016

34. 34. 34

    . Tiznit, Mirght, slopes above the road to Guelmin SW of the village, 910 m, 29º 24.545’N, 9º 43.574’W, rocky and bushy slopes dominated by Euphorbia, 14-III-2015

35. 35. 35

    . Assa,Targoumait, along the road to Assa 15 km SE of the town, 500 m, 28º 46’ 32.27”N, 9º 27’ 29.84”W, rocky and sandy plane with solitary Acacia trees, 15-III-2015

36. 36. 36

    . Assa, edge of dry riverbed at SW edge of the town, 310 m, 28º 36’ 8.02”N, 9º 26’ 39.99”W, Acacia trees and bushes on dry rocky slope, 15-III-2015, 24-III-2017

37. 37. 37

    . Assa, in the oasis at the town centre, 300 m, 28º 36’ 19.93”N, 9º 25’ 40.04”W, abandoned meadows, dry riverbed, 15-III-2015

38. 38. 38

    . Assa, small wadi at the turn for Tuisgui Remz Jdid from the road to Zag, 290 m, 28º 26’ 46.37”N, 9º 21’ 56.33”W, dry riverbed with Acacia trees, 15-III-2015, 21-III-2016, 24-III- 2017

39. 39. 39

    . Assa, Tistguezzemtz, oasis at the dam in the village, 230 m, 28º 24’ 53.86”N, 9º 13’ 1.21”W, bushy road verges, rocky slopes, ruderal areas, 15-III-2015, 24-III-2017

40. 40. 0

    4. Assa, large wadi south of Ouarkziz Mts. Along the road to Zag, 250 m, 28º 22’ 45.31”N, 9º 23’ 11.52”W, bushy and sandy area with Acacia, 15-III-2015

Results

Discussion

A total of 54 species were encountered during our surveys representing roughly 40% of the fauna of Morocco. The number of observed species exactly matches the number of species listed for Anti- Atlas (excluding Djebel Sioura) by TARRIER & DELACRE (2008), however the skippers are not included in their list. Given the fact that our visits were limited to March and April, thus excluding most of the late flying satyrids, the number of encountered species could be considered exceptionally high.

Among the most common and widespread species encountered during our surveys are hábitat generalists like Pieris rapae, Pontia daplidice, Euchloe charlonia, Euchloe crameri, and Lycaena phlaeas which thrieve in cultivated landscapes. Vanessa cardui was common and widespread only in years with abundant precipitation (like in 2009) when the region becomes an important source area for its migration to Europe (STEFANESCU et al., 2011). On the other hand, several rare species were observed and these are briefly discussed below:

• Papilio saharae (Fig. 7).- This is a known eremic species limited to arid regions at the northern edge of the Sahara and western part of the Arabian Peninsula (PITTAWAY et al., 1994). Two specimens were observed along a ridge south of the Col du Kerdous in mid March, so well outside the flight period of the more widespread sister species P. machaon (TARRIER & DELACRE, 2008). One of the specimens was already extremely worn indicating its posible emergence as early as February.

• Zerynthia rumina.- According to TARRIER & DELACRE (2008) the species is widespread in Djebel Lekst area near Tafraoute, but we were unable to find any despite targeted searching. Our only observation is from the Tiouite village oasis just south of the Souss Valley.

• Euchloe falloui (Fig. 4).- Another eremic species limited to desert regions of North Africa and Arabian Peninsula (TENNENT, 1996). Although considered rare in Morocco, the species was widespread and common in 2015 following the rainy winter in the Anti-Atlas, in some places outnumbering the similar and more widespread E. belemia. A similar phenomenon was also observed by Tennent in 1994 (TENNENT, 1996). Our records are from Assa region in the south and at much higher altitudes to the east of Tafraoute and Igherm.

• Colotis chrysonome (Fig. 6).- There is very limited information available on the distribution of this conspicuous pierid in Morocco where it is considered migratory (TENNENT, 1996). Tarrier’s record from the north-eastern Anti-Atlas at Tazenakht falls into that category, as it is beyond the range of the species host plant Maerua crassifolia (Capparidaceae). According to RUNGS (1972) the species was found in lower Draa Valley near Auinet Torkoz north-west of Tan Tan. Based on available cartography this town is however much further upstream near Assa giving us a hint to visit this region in search for the species. We found both the host plant and adults in a small wadi south of Assa (Fig. 8) on the road to Zag. Four fresh males were observed in mid March 2015 and in March 2016 the species was again seen in some numbers at the same site. This indicates that the species is resident in this region or at least forms temporary colonies.

• Colotis liagore.- Published here for the first time for Morocco and only previously known from the southernmost part of Algeria in the Maghreb (TENNENT, 1996). It shares its host plant with the previous species, therefore its presence in the Assa region could be anticipated. The two specimens observed at the site south of Assa in 2016 were however still a big surprise. They were syntopic with C. chrysonome. Given the presence of its host plant, it is possible that it builds temporary colonies in this part of Morocco, however further surveys are needed to confirm its continuous presence so far north. It must be noted that in 2010 the species was photographed on the road between Dhakla and Aoussard much further south in the West Sahara territory (Raoul Beunen, pers. comm.), this being the first record of the species for the country.

• Callophrys avis.- This typical west Mediterranean species has so far not been mentioned from the southern Anti-Atlas (TARRIER & DELACRE, 2008). Here we publish records from the region for the first time: the earliest was by Stefano Bossi in 2005 (Bossi, pers. comm.) and we confirmed its presence at Tizi’ N Tarakatin by finding two specimens in a dry wadi north-west of the village in March 2017.

• Cigaritis allardi.- This attractive lycaenid has very restricted colonies and is therefore easily overlooked and probably under-recorded. We found it at two sites on very different biotopes. The site near Djebel Lekst the habitat corresponds well with the description by TENNENT (1996) - steep rocky slopes covered by bushes, whereas the site at Mirght represents a pasture heavily overgrown with cactus like Euphorbia sp. and thorny brooms (possibly Genista sp.), which could be the host plant of the species.

• Tarucus rosaceus.- In contrast to the scattered records for this species indicated in Tarrier & Delacre (2008) it proved to be much more common in the south. We found it both at higher altitudes near Tafraoute where it is sympatric with T. theophrasuts and near Assa where it is the only species of the genus present.

• Azanus ubaldus.- Despite its widespread occurrence in dry tropical regions of Africa and Asia including neighbouring regions (TENNENT, 1996) the species has only recently been confirmed for Morocco (WEISS, 2000), quite unexpectedly for the northern Atlantic coastal region. The species was common on the outskirts of Assa at the edge of a dry wadi during visits in 2015 2016, and 2017. A copula was observed also in a small wadi south of Assa en route to Zag in 2015. Given the abundance of the acacia trees the species is most probably a permanent resident in the region.

• Plebejus allardi (Fig. 5).- First reported for Anti-Atlas by BOZANO & GIACOMAZZO (1988) and then raised to a specific rank (P. antiatlasicus) by Tarrier (1995). Regardless of its status, the taxon has a very restricted range limited to the southern Anti-Atlas Mountains. We found the species at several sites east of Tafraoute and east of Igherm always in the presence of its larval host plant Astragalus caprinus L. which is usually detected before the butterfly. The reported flight period of the species is from April to May (TARRIER & DELACRE, 2008), however we found the species regularly in mid-March, in some cases even worn specimens.

• Melitaea deserticola (Fig. 1).- Due to its similarity to the more widespread M. didyma its distribution in Morocco is poorly understood. Although our findings are congruent regarding distribution patterns of both species, M. deserticola was not rare and was observed syntopic with M. didyma at two sites near Azgour and Taghaout, both east of Tafraoute.

• Spialia doris (Fig. 2).- This is another eremic species. It has a restricted and fragmented distribution in northern Africa, where it is known only from southwestern Morocco and Egypt. We found the species at several dispersed sites in the southern Anti-Atlas at elevations between 1200 and 1600 m. Interestingly we also found it further south at Assa along the road to Zag in much more arid semidesert environments. It is very likely that the species is also present in similar habitats in nearby Algeria from where it has not been recorded.

The butterfly data coverage for the southern Anti-Atlas, and in Morocco in general, is still insufficient and there are many regions with very limited or no butterfly records. A more systematic approach is needed with future surveys targeted at covering at least some of the unstudied areas. Although the grazing pressure is less intense in Anti-Atlas compared to the Middle and parts of the High Atlas (authors, pers. observ.), negative impacts of overgrazing are evident especially around villages and areas with easy access. As butterflies are very sensitive to such anthropogenic pressure (Numa et al. 2016) our data will be of great value for future comparisons.

Acknowledgments

We wish to thank the following people for their company and help during our surveys: Luca Azzolin, Alessandro Floriani, Primoz Glogovgan, Marko Kosmag, Alessandro Maculan, Enrico Mojentale, Peter Rowlings, Luca Sattin, Giuseppe L. Turri, Peter Valiè, and Barbara Zaksek. We are especially grateful to Luca Sattin for the contribution of a superb photograph of Colotis chrysonome and Miguel López Munguira for the translation of the abstract. We also thank Raoul Beunen for sharing his record of Colotis liagore.

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