Crustacea, Amphipoda

Zootaxa 2546: 31–51 (2010) www.mapress.com / zootaxa/

ISSN 1175-5326 (print edition)

Article

Copyright © 2010 · Magnolia Press

ZOOTAXA ISSN 1175-5334 (online edition)

Two new Gammarus species (Crustacea, Amphipoda) from warm springs in the south-east pre-alpine area of the Zagros, Iran: habitats with physiological challenges MEHRDAD ZAMANPOORE1, MICHAL GRABOWSKI2, MANFRED POECKL1 & FRIEDRICH SCHIEMER1 1

Department of Limnology, Institute of Ecology, University of Vienna, Austria. E-mail:[email protected], [email protected], and [email protected] 2 Department of Invertebrate Zoology & Hydrobiology, University of Lodz, Poland. E-mail: [email protected]

Abstract Despite the prospering results emerging from some recent works on Iranian amphipod fauna, continental waters in the south-east edges of the Zagros Mountain ranges had not been investigated for gammarids before. This paper introduces two new epigean species, G. shirazinus and G. loeffleri, distributed in many running waters of four catchment areas in a broad region in the south of Iran. The most important ecological features of the habitats of these species in comparison with previously studied gammarids are higher water temperature and salinity, and lower dissolved oxygen in their habitats, coinciding with decreasing elevation. A comparison of the ultra-structure of the cuticle of the head capsule by scanning electron microscopy between these two new species, and some other species is given. A detailed species description is provided. In addition, the geographical and ecological range of the new species is discussed. Key words: Amphipoda, Gammaridae, Gammarus shirazinus, Gammarus loeffleri, Zagros, Fars, Iran, SEM, water temperature, salinity

Introduction Biodiversity of amphipod crustaceans in the Southern Zagros Region has been investigated in some recent works (Mateus & Mateus, 1990; Stock et al., 1998; Zamanpoore et al., 2009). All these studies introduced several new species to science, as well as those of central Zagros Mountains (Khalaji-Pirbalouty & Sari, 2004; 2006), revealing a high potential of amphipod speciation in the region. Almost all of the analyzed samples, however, were collected from cold springs of mountainous highlands, while a broad region in the south-east skirts of the Zagros mountain-chain was entirely deprived of scientific attention. Southern margins of Zagros are comprised of many of totally or partially isolated catchment areas. It is well known that among several environmental mechanisms involved in isolation of populations in the course of evolving new taxa, geographical isolation by river catchments is of special importance for aquatic life (Banarescu, 1990). On the other hand, some ecological factors like water temperature and salinity shift to much higher levels as habitats lose altitude at the mountains' peripheries. This paper presents results of examination of samples from fresh water springs and rivers in various catchment areas in southern margins of the Zagros Mountains, Fars Province, Iran. Two new species are described in detail using macro-morphology and cuticular micro-structures, with an emphasis on the usefulness of scanning electron microscopy (SEM) in amphipod taxonomy. Some ecological data with important variations to the habitats of previously established species are presented and their possible roles are discussed.

Accepted by G. Karaman: 5 May 2010; published: 23 Jul. 2010

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Methods Samples were taken from four catchment areas in Fars Province, Iran (Fig. 1), two times in February and August 2007 for type localities; other sites have been sampled only once or twice at various dates since 1995. Specimens were collected using a hand sieve from beneath the leaves of aquatic plants, from gravel banks (after agitating bed) or from autumn-shed, decaying leaf litter accumulated by debris dams in their streambed, according to the natural stream habitats. They were fixed in 75% ethanol in locus, followed by three weekly replacements afterwards in the lab. Descriptions were made based on the holotype and additionally 5 winter and 5 summer paratypes from the type localities; these paratypes were dissected and parts were stained with Lignin Pink (Sigma Aldrich), mounted using Euparal (Carl Roth GmbH & Co) on microscope slides. A complete series of digital microphotographs was taken from body parts after dissection, which was later used to make digital drawings in CorelDRAW (V.11.633, 2002, Corel Corporation).

FIGURE 1. Map of the study area from Central to Southeast of Fars Province, Iran. Bold lines indicate borders of river catchments.

Five adult individuals from each of the two species (from the type locality) were prepared in the next step for producing scanning electron micrographs, following the methods originally described by Haley (1997). Various regions on laterals of the head capsule were scanned and photographed through a serial magnification. Some materials from the region, previously collected and deposited in the Natural History Museum of Vienna (NHMW) and the Zoological Museum of the University of Amsterdam (ZMA) were also examined. These include G. crinicaudatus Stock et al., 1998 (holotype, ZMA Crust amph 201937), G. komareki Schaeferna, 1922 (paratypes from Shahrestanak, Iran; ZMA Crust amph), and Gammarus syriacus Chevreux, 1895 (material from NW Shiraz, Iran; NHMW, 4867). Ecological data were measured on the same day from 0900 to 1100. Dissolved oxygen and pH were measured by a WTW-Oxy 320 and a HANA-HI 1281 probe, respectively, at their microhabitat – restricted to small areas where amphipods were caught. A volume of 100 ml of filtered (0.45 μm Whatman paper) spring water kept in cold box were used for the measurement of ionic content.

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Results Gammarus shirazinus n. sp. (Figs. 2–5) Gammarus syriacus Chevreux, 1895, [in Mateus & Mateus 1990: 280].

Material Examined. Many males and females. Male holotype, body length 22 mm, February and August 2007, coll. M. Zamanpoore. Locus typicus: Pole-Berenji spring, 10 km S of Shiraz, Fars Province, Iran (2927Ń, 5231É), Altitude 1490 m. Paratypes, many males and females; same date and locality. Additional samples: Barme-Delak spring, 18 km SE of Shiraz, (2933Ń, 5242É), Altitude 1467 m.; Barme-shur spring, 17 km S of Shiraz, (2928Ń, 5241É), Altitude 1463 m.; Barme-Tarkoshte spring, 17 km SE of Shiraz, (2934Ń, 5240É), Altitude 1472 m.; Kaftarak, 16 km SE of Shiraz, (2935Ń, 5239É), Altitude 1473 m.; Niriz, 200 km SE of Shiraz, (2912Ń, 5420É), Altitude 1603 m.; Pire Bano spring, 11 km SW of Shiraz (2931Ń, 5227É), Altitude 1492 m.; Pire Gheibi spring, 12 km SW of Shiraz (2931Ń, 5227É), Altitude 1492 m.; Se Barm spring, 14 km E of Shiraz (2935Ń, 5240É), Altitude 1488 m.; All specimens including type series are kept in the amphipod collection of the Museum of Fars Research Centre of Agriculture and Natural Resources (FARSAGRES), Shiraz, Iran. Holotype is deposited in the Zoological Museum of the University of Amsterdam (ZMA, Amph. 206057, 2008). Diagnosis. A medium species. In general, this species can be distinguished from other adjacent species by a combination of the following characters: (1) highly setose antenna 2 and pereopods 5–7 posterior margin, (2) antenna 1 with two spines on the ventro-distal of first peduncle, (3) no setae on postero-distal corner of basis pereopod 6 and 7, and (4) longer fine setae on posterior pereopods 5–7 basis. Etymology. The species' name shirazinus is made from the name of Shiraz, Capital of Fars Province. The type locality is in a very close distance southwards. Description. Description is based on examination of the holotype, 10 male and one female paratypes. Male. Maximum body length (based on 40 individuals) 23 mm.; lateral cephalic lobes rounded, anterior lower part of the head not extended nor elongated; eyes reniform, of the medium size (the same length as the diameter of the first segment of antenna 1, length of which twice or less than twice as wide (Fig. 2D). Antenna 1: long, about half the body length; peduncle segment 3 less than half the length of the first and the second segment; a group of some 3–5 setae on lateral sides of the first peduncle tip, and two spines on its ventral side (Fig. 2B); 2–3 groups of short setae (the same length as the width of the related segment) on ventral of the second peduncle segment and one group on the third; main flagellum with 30–36 and accessory flagellum with 3 segments, respectively, with very short simple setae in groups of 3–5 on each side (Fig. 2A). Antenna 2: Gland cone do not reach the distal end of the third peduncle segment; peduncle segments 4 and 5 of the same length, with groups of long setae (up to three times the length of the diameter of the segments); flagellum with 10–16 segments; long setae, the longest of which are set at the proximal segments, groups of setae sub-marginal to the tip of each flagellar segment at both outer and inner surfaces, a row of setae sub-marginal to the ventro-distal surface of each segment, flag-like brush not formed; calceoli present (Fig. 2C). Mandible: With well developed incisor processes and lacinia mobilis, a plumose long spine row and a ridged molar process (Fig. 2E). First segment of mandible palp not armed; ventral setae on the second segment comprised of 3–4 separated short setae (shorter than the width of the segment) at the proximal and 6 longer setae (as long as to slightly longer than-three times the width of the segment) implanted very close to each other at the distal; a comb-like row of 28–32 D-setae, 5 long E-setae, a groups of B-setae (5) and a group of A-setae (3) on the external surface of the third segment (Fig. 2F). Labium: Simple; a group of short setae on the inner margin of the apices, a group of bristles on more proximal of the apex of the inner lobe, and a group of very fine bristles on the outer margin. Maxilla 1: long plumose setae on inner lobe; outer lobe with stout serrate spines; apex of the left palp with 6 median spines accompanied by 3 median setae (2 times longer than spines), one longer separate subapical spine on distal outer corner (Fig. 2H), right palp with six robust tooth-like spines on the apex, one TWO NEW GAMMARUS SPECIES FROM SOUTHERN ZAGROS

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longer separate spine in a sub-apical position on its distal outer corner and one longer setae between them (Fig. 2I).

FIGURE 2. Gammarus shirazinus n. sp., holotype, %, 22 mm., from Pole-Berenji spring, S of Shiraz. A: antenna 1, B: antenna 1, enlarged ventral tip of first peduncle, C; antenna 2, D: head capsule, E: mandible, F :Palp of mandible, G: exopodite of Maxilipede, H: left palp of maxilla 1, I: right palp of maxilla 1, and J: telson.

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FIGURE 3. Gammarus shirazinus n. sp., holotype, %, 22 mm., from Pole-Berenji spring, S of Shiraz. A: gnathopod 1, B: gnathopod 2, C: pereopod 3, and D: pereopod 4.

TWO NEW GAMMARUS SPECIES FROM SOUTHERN ZAGROS


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Maxilliped: Distal margin of exopodite with a row of three strong teeth and 8 longer setae; a single spine proximal to the most interior teeth, a row of three setae parallel to the long axis close to the single spine, a row of setae extended from proximal to distal on interior margin, shorten gradually, becomes sub-marginal under the distal apex, (Fig. 2G); endopodite with a row of long plumose setae distally, losing plumosity and shortening interiorly, merging with a row of inferior spine-teeth proximally, which gradually shorten; palp well developed. Gnathopod 1: Distal of coxal plate with the same width as proximal, corners rounded, one seta at postero-distal and 1–2 setae at antero-distal corner, 1–2 longer setae at distal sub-marginal surface of the plate; basis slightly expanded at 1/3rd, getting narrower proximally; posterior margin of carpus and merus with groups of setae which are plumose on carpus; propodus pyriform, palm oblique with both medial palmar and palmar angle spines, propodus with 4–5 groups of small spines at posterior margin and 2 groups of longer setae (2–3 each) on posterior sub-margin, dactylus long, a group of four setae very close to medial palmar spine, two groups of shorter setae at anterior margin, 2 rows of short setae (5 groups each) on propodus face (Fig. 3A). Gnathopod 2: Distal part of coxal plate with almost the same width as the proximal part, one seta at postero-distal and 2 setae at antero-distal corner, 2–3 setae on the distal sub-margin of the plate surface; propodus sub-rectangular, more elongated than propodus of gnathopod 1; palm oblique with both robust medial palmar and palmar angle spines (2), 3–4 additional small spines close to the palmar angle spines; a group of long setae attached to the base of medial palmar spine, two rows of long setae (of 4–5 groups each) on the propodal surface, but no groups on anterior margin (Fig. 3B). Pereopod 3: Coxal plate rounded at antero-distal corner, posteriorly concave and anteriorly convex margin, with a setae at postero-distal corner, and a longer setae on the surface of the plate; posterior margin of basis-carpus with groups of setae, length of which about 1 to 1.5 times the diameter of the segments in basis and ischium; in merus and carpus they are in dense groups, 2–3 times longer than the diameter of the segments; merus anterior margin with 2–3 groups of few setae and a spine, a group of long setae and a spine at anterior tip; carpus posterior margin with one spine among some of its 8–9 groups of setae, and a long spine and a group of longer setae implanted on both its anterior and posterior tip; 5 groups of one spine and many long setae on propodal posterior margin (Fig. 3C). Pereopod 4: The lengths of setae are shorter, and the number of setae and groups of setae are lower than those in pereopod 3. Two small setae on antero-distal and 4 at postero-distal margin of coxal plate; merus anterior margin with two groups of short setae and one spine, one long spine among a group of setae implanted at anterior tip; carpus with two spines in some groups of posterior margin armature (Fig. 3D). Pereopod 5: Basis sub-rectangular, protruding lobe developed at postero-distal, about 10 setae at posterior margin which are long for this place (as long as or longer than the length of ischium), anterior basis with a marginal row of 1–2 groups of setae (2–3 each) and 3–4 spines; small spines and longer setae as long as the segment width on merus, but more than 2 times longer than the segment width on carpus, propodus with 5 rows of spines (3–3–3–3–2) on anterior margin and long setae on posterior (Fig. 4A). Pereopod 6: Longer than pereopod 5; basis more slender, postero-distal protruding lobe weakly developed; setae on all segments much longer than those of pereopod 5, nearly 3 times the length of the related segments; 6 rows of spines on propodus anterior margin (Fig. 4B). Pereopod 7: Basis wider proximally than distally; postero-distal lobe weakly developed, as an obtuse corner, some 15 setae on posterior margin, slightly shorter than the length of ischium; merus and carpus anterior margin with long spines and very long setae (up to 4 times the width of the related segment), posterior margin with less numerous and shorter setae; propodus with shorter spines (2–3–3–3–3–2) and setae on anterior margin, and setae only on posterior; posterior tip of propodus with 2 spines mixed with a group of longer setae (Fig. 4C). Uropod 3: Endopodite about 3/4 (Mean = 0.73, SD = 0.02) of the exopodite; setae on both rami long but shorter on proximal part of exopodite, many plumose setae on outer margin of both exo- and endopodite, three groups of long robust spines sometimes with few setae implanted on tip of basis; external margin of exo- and endopodite armed with long spines implanted in intervals (Fig. 4F).

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FIGURE 4. Gammarus shirazinus n. sp., holotype, %, 22 mm., from Pole-Berenji spring, S of Shiraz. A: pereopod 5, B: pereopod 6, C: pereopod 7, D: urosomites, E: epimeral plates, and F: uropod 3.



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FIGURE 5. Gammarus shirazinus n. sp., paratype, &, 12 mm., from Pole-Berenji spring, S of Shiraz. A: basis of pereopod 5, B: basis of pereopod 6, C: basis of pereopod 7, D: epimeral plates, and E: oostegite 1.

Epimeral plates: Epimeral plate 1 rounded with a tiny pointed node at postero-inferior corner, some 14 setae on antero-inferior corner; anterior margin of the second plate rounded and slightly pointed at posterior corner; third plate moderately pointed; epimeral plates 2 and 3 with 1–2 spines and a few longer setae on inferior sub-margin, one-two long setules on distal posterior margin in all 3 plates, posterior dorsal margin of metasomites 2 and 3 with a row of long setules (Fig. 4D). Telson: Elongated, lobes about twice longer than broad; a group of two–three long and robust spines and some 7 long setae on the tip; dorsal surface of the lobes with 3–4 groups (of 2–4) setae (Fig. 1J). Urosomites: posterior dorsal surface of urosome segments without elevation, dorsal armature of 1–2 spines on latero-dorsals and 2 spines on mid-dorsal, mixed with some short setae (Fig. 4E). Female. The female is smaller than male (12 mm). Palm less oblique in propodus of gnathopod 1; no medial palmar spine, a long palmar angle spine, 2–3 spines at posterior palmar margin, and a group of setae at medial palm. Propodus of gnathopod 2 sub-rectangular, without the medial palmar spine; 3 palmar angle spines, a group of setae at medial palm, posterior palmar margin without spines. Propodus of gnathopod 1 and gnathopod 2 is smaller than in males, compared to the other segments and the whole appendage than in males. Proximal 3/4th in bases of gnathopod 1 and gnathopod 2 are not wider than their distal. Bases of pereopods 5–7 are less elongated compared to males, with distinctly convex angles at the middle of posterior margin (Fig. 5A–C); shorter setae on distal segments. Epimeral plates 2 and 3 without setae on distal sub-margin, showing only few spines (Fig. 5D). Oostegite 1 is illustrated in Figure 5E.

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Cuticular ultrastructure. Cuticular polygons of head capsule show both linear (L-type) and scattered (Stype) types of pore arrangements (Fig. 6A); distances between lines were 0.97–2.22 (Mean = 1.56 , SD = 0.24); density of pores in L-type polygons were 54–75 pores per 25 µm2 (Mean = 64, SD = 5.04) while S-type polygons had higher densities of 104–170 pores per 25 m2 (Mean = 131, SD = 14.72), arranged in a specific cluster-shaped pattern (Fig. 6C).

FIGURE 6. Micrographs of ultrastructural patterns on head region in Gammarus shirazinus n. sp. (A & C), and G. loeffleri n. sp. (B & D).

Remarks. Despite of some similarities among all populations of the whole area, one may not encounter with much difficulty in identifying G. shirazinus from other adjacent species, using the clear distinctive feature of having 2 spines on the first peduncle of antenna 2. Comparisons of the new species with other closely related species are given below. Mateus and Mateus (1990) examined samples from the very same area (material from Crustacean Collection of the Natural History Museum of Vienna (NHMW), coll. no. 4867, one male, one female) identified them as Gammarus syriacus Chevreux, 1895. Re-examination of these samples showed that they are completely identical to our samples, especially in having the antenna 1 spines. Here is a comparison of the description of G. syriacus from Syria and Lebanon given in Karaman and Pinkster (1977, pp. 60–64, Figs. 24A–L; 25A–L) with our newly described species. In G. syriacus antenna 2 peduncles and flagellum bear short setae, and the gland cone is only half of the third segment, pereopod 3–7 have shorter setae on all segments, pereopod 7 posterior distal protruding lobe is less developed, epimeral plates 1–3 postero-inferior corners are elongated (sabre shaped) and sharply pointed, dorsal humps are quite well developed, and telson is elongate with much short setae. In G. shirazinus all these characters are clearly different. Hence, it seems quite evident that these populations are not con-specific with G. syriacus, and we are dealing with two distinctly separate species. TWO NEW GAMMARUS SPECIES FROM SOUTHERN ZAGROS


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Gammarus shirazinus lives in a region close to localities of G. crinicaudatus Stock et al., 1998 (holotype, ZMA Crust amph 201937). The new species differs from it by having 2 spines on ventro-distal margin of antenna 1 first peduncle segment, longer gland cone (ca. 2/3rd the length of the third peduncle segment) and setae on both peduncle and flagellum and having calceoli on antenna 2, no setae on postero-distal corner of pereopod 7 basis, much longer fine setae on posterior margin of bases and longer setae and spines on distal segments of pereopods 5–7, longer setae on inferior margin of epimeral plates 2–3, telson (more than the length of telson itself), and posterior dorsal margin of urosome. There is also no distinct space between proximal and distal setae on the second segment of palp of mandible, and 28–32 D-setae on inferior margin of the third in the new species, while a clear empty distance and about 20 D-setae are seen in G. crinicaudatus. G. komareki Schaeferna, 1922, widely distributed in the country, shares with G. shirazinus the feature of having a long antenna 1 appendage, long antenna 2 setation, long setae on pereopod 3, similar shape of basis in pereopods 5–7, and long uropod 3 endopodite. On the other hand, it has small eyes, no spines on antenna 1, short gland cone, “very densely setose” with curled setae and slightly swollen and dorso-ventrally compressed antenna 2 flagellum, more numerous distal long setae on second segment and more (40) D-setae on the third segment of palp of mandible, less setose pereopod 4 and pereopods 5–7 distal segments, shorter setae on inferior margin of epimeral plates 2–3, less setation on uropod 3, and much shorter setae on telson, all of which make it totally different from this new species. The only reported species with a similar feature to the two spines on peduncle antenna 1 in all fresh water Gammarus species is G. bakhteyaricus Khalaji-Pirbalouty & Sari, 2004, with three structures on the same place on antenna 1, unless those are called as “short robust setae” by the authors; however we suggest them as the same structure, well recognised as true “spines”, an absolutely rare and unique feature in fresh water gammarids. Despite this similarity, the two species has many differences, among them are small eyes, poorly setose antenna 2, different pattern of setation in gnathopod 1 and gnathopod 2, poorly setose pereopods 5–7 and telson, elongated and tapered pereopod 7 basis with some setae on postero-distal corner, elevated dorsal urosomites, and very short setae on inferior margin of epimeral plates 2–3 (Khalaji-Pirbalouty & Sari, 2004; Figs. 5–7). G. qiani Hou & Li, 2002, described from Yunnan Province, China, has the similarity of setose impression in pereopods 5–7, resembling it to G. shirazinus, but shows many characters differentiating it, including weekly setose peduncles of antenna 1, moderately setose peduncles and flagellum of antenna 2, longer antennal gland cone reaching to the tip of the next peduncle segment, only moderately setose pereopod 3–7, not developed postero-inferior lobe and having corner setae in pereopod 7, only few short setae on an incompletely cleft telson, epimeral plates not pointed, and poorly setose entire uropod 3 with an endopodite which is only slightly shorter than exopodite (Hou & Li 2002; Figs. 1–5). G. translucidus Hou et al., 2004, (from Guizhou Province, China) is also similar regarding the high setosity of the peduncle 4 and 5 of antenna 2 and pereopods 5–7 as well, short antennal gland cone, and poorly developed protruding lobe in pereopod 7, but a cave-dwelling amphipod, it has some important differences with G. shirazinus including lacking the eye, short setae on pereopod 4, setae on postero-inferior corner of pereopod 7 basis, short setae on the telson, and incomplete cleft telson, according to Hou et al. (2004), figs. 1– 4. Ecology. Site description. This new species inhabits several springs and rivers including the locus typicus (Pol'e Berenji) which lies at the bottom of a small chain of hills (1500–2100 m above sea level) at the very beginning of first order brooks in a region very close (10 km aerial and 22 km land distance) to Shiraz, capital of Fars Province in southward direction. The shallow (20–40 cm) spring merges shortly (100 m) with a deeper stream running towards a plain, where it is used up almost entirely in farms during the spring and summer. Other loci, not too far from here, are shown in Figure 1. The spring bed constitutes small sized gravel (Mean diameter = 11.5 mm) and fine sand at the base. Aquatic flora is restricted to Ceratophyllum demersum L. and Nasturtium officinale L., poorly inhabiting the spring, but in much dense populations in the continuing river. Few old trees were previously grown in the margins of the brook, making the desired shadow for gammarids on hot summer days. Unfortunately, all of them were cut shortly before our last sampling in summer 2007 by the nearby village farmers who - in hope of increasing water discharge - have dug the spring canal.

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Ecological data. Mean values for width and depth were 300 cm (SD = 55) and 19 cm (SD = 9) in winter, and 100 cm (almost uniform) and 25 cm (SD = 12) in summer. Water current velocity and discharge were 0.3 m/s and 27dm3/s, respectively, in February, while there was no current/discharge in August. Winter and summer records for water temperature were 21.6°C and 22.4°C, and for dissolved oxygen were 6.5 and 4.4 mg/dm3, respectively at the spring source. Air temperature raised from 15°C to 36°C at the same time, while its absolute maximum records in last 37 years were measured up to 43°C (FRMD 2009a). Other environmental factors measured in winter and summer in the spring are summarized in table 1. Details of some ecological data for other localities are presented in Table 2. TABLE 1. Some environmental parameters in type locality habitats of G. shirazinus n. sp. and G. loeffleri n. sp. G. shirazinus

G. loeffleri

winter

summer

winter

summer

7.3

7.5

7.8

8.0

alkalinity (mval/dm )

3.26

3.28

4.02

3.96

electrical conductivity (µS/cm)

1686

1920

575

564

[Na] (mg/dm3)

213.8

251.8

30.7

30.1

[K] (mg/dm3)

3.3

2.9

1.8

1.5

[Ca] (mg/dm )

75.0

77.3

65.8

65.8

3

[Mg] (mg/dm )

35.3

38.4

15.0

15.5

[Cl] (mg/dm3)

335.2

426.5

37.6

35.6

([SO42-] (mg/dm3)

119.3

126.9

36.5

36.0

Salinity (sum of ions)

781.9

923.8

187.4

184.5

pH 3

3

Gammarus loeffleri n. sp. (Figs. 7–10) Material examined. Many males and females. Male holotype, body length 12.5 mm., February and August 2007, coll. M. Zamanpoore. Locus typicus: Cheshme Golabi spring, 23 km W of Darab, Fars Province, Iran (28˚47′ N, 54˚22′ E), Alt. 1100 m. Paratypes, many males and females; same date and locality. Additional samples: Aab Do-man-nim spring, 12 km S of Kherame, (28˚42′N, 54˚27′E), Alt. 1090 m.; Aab Vohli spring, 22 km S of Kherame, (28˚42′N, 54˚27′E), Alt. 1090 m.; Atashkade spring, 13 km N of Firooz Abad, (28˚53′N, 52˚32′E), Alt. 1366 m.; Bahadoran river, 15 km W of Darab, (28˚42′N, 54˚27′E), Alt. 1090 m.; a small river in Bande Amir road, 41 km E of Shiraz, (29˚33′N, 52˚58′E), Alt. 1570 m.; Fasa river, 5 km N of Fasa, (29˚01′N, 53˚38 ′ E), Alt. 1430 m.; Ghare Aghaj river, 53 km S of Shiraz, (29˚11 ′ N, 52˚38 ′ E), Alt. 1650 m.; Nowbandegan, 20 km SE of Fasa, (28˚50′N, 53˚49′E), Alt. 1250 m.; Oghlanghez spring, 30 km NW of Darab, (24˚50′N, 54˚24′E), Alt. 1540 m.; Pa-Naal spring, 10 km NW of Fasa, (29˚07′N, 53˚27′E), Alt. 1520 m.; Sarve Nokhodi, 105 km SE of Shiraz, (29˚11′N, 53˚06′E), Alt. 1490 m.; Tang Aab spring, 29 km N of Firooz Abad, (29˚02 ′ N, 52˚34 ′ E), Alt. 1540 m. Type series are kept in the amphipod collection of the Museum of FARSAGRES, Shiraz, Iran. Holotype is deposited in the Zoological Museum of the University of Amsterdam (ZMA, Amph. 206055, 2008). Distribution of the species is shown in Figure 1. Diagnosis. A small species with elongated eyes; this species can be distinguished from other similar species by combination of these characters: (1) highly setose antenna 2, (2) pereopod 7 basis without protruding lobe but with setae on postero-distal corner, (3) pointed postero-distal corner of epimeral plates 1– 3, (4) long setae on dorsal of urosome, (5) long setae on anterior margin of pereopods 5–7, (6) long spines mixed with setae on anterior and posterior of carpus and merus in pereopods 5–7, (7) and long setae on telson apex and surface.



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FIGURE 7. Gammarus loeffleri n. sp., holotype, %, 12.5 mm., from Cheshme Golabi spring, W of Darab. A: antenna 1, B: antenna 2, C: head capsule, D: mandible, E: palp of mandible, F: exopodite of maxilliped, G–H: left palp of maxilla 1, I: right palp of maxilla 1, and J: telson.

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FIGURE 8. Gammarus loeffleri n. sp., holotype, %, 12.5 mm., from Cheshme Golabi spring, W of Darab. A: gnathopod 1, B: gnathopod 2, C: pereopod 3 and D: pereopod 4.



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Etymology. We take honour in naming this species for the late Professor Heinz Loeffler of the University of Vienna, in gratitude of his founder contribution to Iranian limnology, especially his studies on Lake Niriz, which is in the same geographical area of the present work. The whole study was planned to be conducted under his supervision, before he passed away in 2006. Description. The description is based on the examination of the holotype, 10 male and one female paratypes. Male. Maximum body length in 18 samples 14 mm. Lateral cephalic lobes more or less rounded, anterior lower extension of the head weekly developed; eyes elongated (longer than the diameter of the antenna 1), twice as long as wide, and reniform (Fig. 7C). Antenna 1: generally long, about 2/3rd–3/4th the total body length; third peduncle segment is slightly shorter than the first and the second; groups of setae on lateral mid-length of peduncle segments 2 and 3; main flagellum with 27–30 segments with very short setae; accessory flagellum with 3–4 segments (Fig. 7A). Antenna 2: Gland cone short, reaching to ca. the half of the length of the third peduncle segment; peduncle segments 4 and 5 with the same lengths, with dense groups of long setae (more than twice the length of the diameter of the segments); flagellum with 11–16 segments, not swollen nor compressed; groups of setae on the tip of the flagellum segments at dorsal and lateral faces, additional row of setae implanted submarginal to the distal of each segment at its ventral surface, setae on flagellum are shorter than peduncle segments and do not form a flag-like brush; three to six calceoli from segment 2 on (Fig. 7B). Mandible: Incisor processes and lacinia mobilis well developed, in addition of a plumose long spine row and a ridged molar process (Fig. 7D). In mandible palp, ventral setation of the second segment include ventral setae, 3–5 shorter (as long as or a little longer than the width of the segment) single setae at proximal and 4–6 longer (longer than twice the width of the segment) setae in a row at distal; inferior margin of the third segment armed with a comb-like row of 18–20 D-setae, 5 long E-setae, one group of A- and one group of Bsetae (Fig. 7E). Labium: Simple, with a group of short setae on the inner lobe apex, and a group of very fine bristles on inner and outer lobes. Maxilla 1: Inner lobe with long setae having fine plumes; 8–10 stout serrate spines on outer lobe; right palp with five to six short and stout spines on the apex, one longer separate spine on its distal outer corner and a setae near it (Fig. 7I); on the apex of the left palp is an armature of four to five spines and a longer setae (Fig. 7H); variations in the strength of spines in some individuals (Fig. 7G). Maxilliped: Exopodite having three–four strong teeth and three–four longer setae at the distal margin; a row of long setae at distal sub-margin which becomes plumose from the middle and continues towards the inferior margin to join to 6–7 very long plumose setae, a strong spine with a distance at sub-marginal interior corner (Fig. 7F); endopodite has a row of strong plumose setae on distal margin which continues to the half of the interior margin, while the setae gradually shorten, get stronger spine-shaped, and lose plumosity; a row of long setae on the whole of interior sub-margin shortening proximally, a row of very fine and long setae on the whole of exterior margin; the pulp is well developed. Gnathopod 1: Coxal plate rectangular with rounded corners, antero-distal corner with 3–5 and posterodistal corner with one short setae, 2–4 longer setae on the postero-distal surface; distal 2/3rd of basis is wider, slightly expanded at 1/3rd, gradually gets narrower proximally; carpus posterior margin with groups of strong plumose setae; propodus pyriform, palm oblique, having both medial palmar and palmar angle spines, posterior propodus margin with a row of 3–5 groups (1–2 each) of spines, continuing proximally with one– two groups of long setae, 1–2 groups of very short spines (1–2 each) on posterior propodus sub-margin, 2–3 groups of fine short setae comes behind them proximally; 2–3 groups of setae at propodus anterior margin; dactylus is long, a group of four long setae very close to medial palmar spine (Fig. 8A). Gnathopod 2: Distal of coxal plate slightly narrower than proximal, anterior distal corner with 3–4 and posterior corner with one short setae, 2–3 longer setae on the surface in posterior distal position; posterior margin of propodus is straight so that makes a rectangular shape, palm is more or less oblique with a robust medial palmar and 3 palmar angle spines; a group of setae beneath the medial palmar spine, 2–3 groups of setae at propodus anterior margin (Fig. 8B).

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FIGURE 9. Gammarus loeffleri n. sp., holotype, %, 12.5 mm., from Cheshme Golabi spring, W of Darab. A: pereopod 5, B: pereopod 6, C: pereopod 7, D: urosomites, E: epimeral plates, and F: uropod 3.



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FIGURE 10. Gammarus loeffleri n. sp., holotype, &, 7.5 mm., from Cheshme Golabi spring, W of Darab. A: basis of pereopod 5, B: basis of pereopod 6, C: basis of pereopod 7, D: epimeral plates, and E: oostegite 1.

Pereopod 3: All segments bear groups of setae at posterior margin, about 1–2 times as long as the diameter of the segments, those of merus and carpus are in dense groups; merus on anterior margin with two groups of mixed setae and one spine, and a same group at anterior tip; anterior margin of carpus with a spine among first three proximal groups of setae, a long spine and a group of longer setae at anterior tip and the same but with two long spines at posterior tip; anterior margin of propodus with 5 groups of one small spine and some longer setae, the last of which does not reach to the anterior tip; coxal plate rounded at distal, concave at posterior, and convex at anterior margin, four small setae implanted at distal anterior and one at distal posterior margin (Fig. 8C). Pereopod 4: Similar to pereopod 3, but the basis is wider; merus on anterior margin with a group of two short setae and two shorter spines on the middle of the segment, and another group of long setae mixed with two long spines at anterior tip; carpus with 2 spines and a group of long setae at posterior tip; anterior margin of propodus with 4 groups of one small spine and some longer setae; coxal plate expanded at posterior distal half, with 4 small setae implanted at distal anterior and 7 at distal posterior corner (Fig. 8D). Pereopod 5: Basis sub-rectangular and slightly concave at posterior margin, backward protruding lobe at posterior distal developed, posterior margin with 13 very short setae, a row of 2–4 setae and 3–4 spines mixed with a very fine seta at anterior margin; merus and carpus anterior margin with long spines and setae (longer than spines, about 1–2 times than the segment’s width), with shorter and less dense setae at posterior; anterior margin of propodus with 6–7 rows of spines ((1–)2–3–3–3–3–2) mixed with setae, and 5 rows of longer setae at posterior (Fig. 9A). Pereopod 6: Similar to pereopod 5, but longer, with longer setae on propodus and carpus anterior and posterior margin; basis is more slender and posterior proximal is slightly expanded, postero-inferior protruding lobe poorly developed (Fig. 9B).

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Pereopod 7: Basis wider proximally; protruding lobe not developed and the postero-inferior corner is set with 1–3 setae, 14 small setae on posterior margin, starting from the very base of the basis; anterior of merus and carpus with long spines and much longer setae; merus and carpus posterior margin with 2 groups (1–2 each) of spine(s) mixed with some shorter setae; propodus anterior with 6 rows of spines (1–2–2–2–2–2) mixed with longer setae, posterior margin with 4 groups of long setae; few long spines at posterior tip of propodus along with a group of longer setae (Fig. 9C). Uropod 3: Endopodite less than 2/3 of the exopodite (ratio of endo-/exopodite length ca. 0.6); very long spines on the distal margin of basis, few setae and 0–1 spine on the basis body; exopodite with long spines at distances on external margin; endopodite with 1–2 spines on external margin; long setae on both external and internal margins of endo- and exopodite, many of which on outer margins are plumose (Fig. 9F). Epimeral plates: Posterior-inferior corner of epimeral plate 1 clearly pointed, 6–8 long setae on anterior distal corner; second and the third plate more sharply pointed; a row of short fine setae on posterior margin of all three plates; a combination of 3–4 setae and spines on distal sub-marginal area of 2nd and 3rd plate (Fig. 9D). Telson: Length of the lobes about twice their widest width; distal margin with 1–2 long and robust spines and some 5–7 long setae; four groups of 3–6 long setae on dorsal surface of the lobes, occasionally accompanied by a long spine (Fig. 7J). Urosomites: Urosome posterior dorsal do not have clear elevations, but each epimere is placed in a lower level than its anterior segment; dorsal armature consists of mainly some 5–6 long setae sometimes mixed with a spine on laterals and mid-dorsal (Fig. 9E). Female. Female is smaller than male (7.5 mm). Antenna 2 with no calceoli. Propodus of gnathopod 1 without medial palmar spine, one long and one short palmar angle spine, 3 spines at posterior palmar margin. Propodus of gnathopod 2 rectangular, palm is transverse and lack the medial palmar spine, but 2 palmar angle spines exist, a group of setae at medial palm. Propodus of gnathopod 1 and 2 is smaller in respect to the other segments and the whole appendage than in males. Proximal of gnathopod 1 and 2 bases are not wider than their distal part. Basis of pereopods 5–7 less elongated than males, rather distinct convex angle at the middle of posterior margin in pereopod 6 and pereopod 7, protruding lobe more developed (Fig. 10A–C). Uropod 3 with plumose setae on outer margin of endopodite only. Epimeral plate 3 with fewer and shorter setae on anterior distal margin, epimeral plates 2 and 3 with 1–2 spines, while no setae on distal sub-margin, (Fig. 10D). Oostegite 1 is illustrated in Figure 10E. Cuticular ultrastructure. Polygons of head capsule cuticles show both types of L and S pore arrangements (Fig. 6B); L-types with line distances of 0.92–2.60 (Mean = 1.79 , SD = 0.24); pore density in L-type polygons 94–123 pores per 25 µ m2 (Mean = 108, SD = 12.45); S-type polygons without a special pattern (Fig. 6D), with density of 143–228 per 25 µm2 (Mean = 176, SD = 36.97). Remarks. There are not many species that this species can be confused with. Some similarities with other species are discussed below. G. loeffleri n. sp., distributed in south-east of the localities of G. shirazinus shows some morphological similarities to it, including long setation of antenna 2 and shapes of pereopod bases and epimeral plates, but it differs from it in many characters, including longer eyes, antenna 1 spines, a shorter antennal gland cone, less groups of setae and spines on gnathopod 1 propodus, less (non-overlapping) number of D-setae on third segment of palp of mandible, non-developed pereopod 7 basis postero-distal lobe, setae on pereopod 7 basis postero-distal corner, shorter very short setae on pereopod 7 basis posterior margin, shorter setae on anterior margin of distal segments especially on pereopod 7, and longer and numerous setae on dorsal of urosomites. G. crinicaudatus Stock et al., 1998, has previously been reported from a close region in the Zagros Mountains. It looks like the new species in having a setiferous antenna 2 and telson, and a similar armature of gnathopod 1. However, there are several characters differentiating G. crinicaudatus from this new species,



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including a shorter eye, no groups of setae on lateral mid-length of peduncle segments 2 and 3 of antenna 1; shorter setae and no calceoli on antenna 2, four small accessory spines close to mid-palmar spine instead of 2 spines close to palmar angle spine, shorter setae and spines on pereopods 5–7, convex pereopod 7 basis posterior margin with more developed posterodistal lobe, weekly pointed postero-distal corner of epimeral plates 2–3, longer spines on the tip of telson, and longer setae on dorsal urosomal armature. G. komareki is widely distributed in northern regions of the country. They have some similarities like an appearance of long setae on antenna 2, shape of basis in pereopods 5–7, and ratio of inner/outer ramus in uropod 3. On the other hand, several features make them distinctly different, among them are long eyes, shorter setae on antenna 2 flagellum, non-curled setae on antenna 2, having calceoli, 19–21 D-setae on palp of mandible, longer setae especially on basis but less dense setae on distal segments of pereopod 3, much more setose pereopod 4, very long setae on propodus-merus anterior margin of pereopods 5–7, having seate on posterior distal corner of pereopod 7, and much longer setae on uropod 3, telson, and posterior dorsal margin of urosome. G. birsteini Karaman & Pinkster, 1977, distributed in Eastern Turkey and Kazakhstan has a similar setose antenna 2, pereopod 7 basis without postero-distal protruding lobe, and the urosome is not elevated. On the other hand, the gland cone in antenna 2 is longer and stays closely to the tip of 3rd segment, setae on pereopods 5–7 are quite short, epimeral plates are not pointed, and, most importantly, inner ramus of uropod 3 is very short and with very few setae (Karaman & Pinkster, 1977; Fig. 31). G. rambouseki S. Karaman, 1931, has more similarities to this species, including the general setiferous impression, especially in antenna 2, pereopods 5–7, and dorsal of epimeres, shape of pereopod 5–6 basis and the setae on the postero-distal corner of pereopod 7 basis, and pointed postero-distal corner of epimeral plates 2–3, but it has clear differences like a small eye, more number of setae on second segment of mandibular palp and a group of A-setae on the third, lack of calceoli, having submarginal spines on propodus of gnathopod 2, and uropod 3 with shorter setae which all are simple (Karaman & Pinkster, 1977; Fig. 32). The setouse character can also be seen in G. frater Karaman and Pinkster, 1977, which shows the similar shape of pereopods 5–7 basis and the postero-distal corner of pereopod 7, and has no elevation in its epimeres, while bears only spines on pereopods 5–7, very shorter setae on pereopod 3–4, telson, and epimeres, and no calceoli on antenna 2 (Karaman & Pinkster, 1977; Fig. 34). Both of these two are specifically local species found exclusively in the Balkans. G. qiani has the similar appearance of setose pereopods 5–7, not-developed pereopod 7 postero-inferior lobe, and the existence of postero-inferior corner setae, but shows actually almost all of its other features different, like a weekly setose antenna 1, moderately setose antenna 2, long gland cone, small eyes, moderately setose pereopod 3–4, short setae on an incompletely cleft telson, not pointed epimeral plates, and poor setation of entire uropod 3, according to Hou and Li (2002), Figures 1–5. Finally, G. translucidus shares some features with G. loeffleri like a moderately setose antenna 1, long setae on fourth and fifth peduncle segment of antenna 2, long setae on pereopods 5–7 (despite they look shorter), and the existance of few setae on pereopod 7 postero-inferior corner. On the other hand, it differs from the new species by having no eyes, longer antennal gland cone, short setae on pereopod 4, slightly developed postero-inferior lobe on pereopod 7, short setae and incomplete cleft on telson, slightly vs. moderately pointed epimeral plates, and scarce short setae on uropod 3 (Hou et al., 2004; Figs. 1–4). Ecology. Site description. This species inhabits a rather large spring-pool (ca. 20 40 m) and the first order stream originating from it located on the bottom of a hill. The stream continues to flow to a plain in southward direction; the water is used up in agriculture in most of the year. This location is 20 km NW of the city of Darab (220 km SE of Shiraz, capital of Fars Province). The spring bed is covered with coarse sand and large gravel, and some organic matter in between them. There are some trees on the stream margins, which amphipods can be most probably found among their free roots in water, as well as inside the decaying leaves trapped in them. There are two pools connected to the stream which are inhabited by some fish populations; amphipods were not found in these pools. Extensive areas at east and west of the spring are devoted to different Citrus orchards.

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Ecological data. This spring is also located in a warm region, where the air temperature reaches to the highest value of some 48˚C in the early afternoon during some summer days (FRMD 2009b). At late morning, we measured the air temperature as 20˚C and 28˚C in winter and summer, respectively, while the measured water temperature had a constant value of 22.5˚C, both in winter and summer. Dissolved oxygen values were recorded at the very low amounts of 3.6 mg/dm3 and 3.2 mg/ dm 3. Other measured physical and chemical environmental parameters are presented in Table 1. Some records of water temperatures measured for additional localities are shown in Table 2. TABLE 2. Records of water temperature (T) in habitats of G. shirazinus n. sp. and G. loeffleri n. sp. G. shirazinus

T (ºC)

G. loeffleri.

T (ºC)

Barme-Delak

17.6

Aab Do-man-nim

15.3

Barme-shur

25

Aab Vohli

20.8

Barme-Tarkoshte

17.6

Atashkade

15

Kaftarak

18.5

Bahadoran river

17.1

Niriz

25

Bande Amir

18.5

Pire Bano

19.7

Fasa river

19.5

Pire Gheibi

20

Ghare Aghaj river

22

Se Barm

21

Nowbandegan

23.4

Oghlanghez

20.8

Pa-Naal

25

Sarve Nokhodi

11.9

Tang Aab

21

Discussion Taxonomy. G. shirazinus has some shared features with G. crinicaudatus, G. komareki, G. bakhteyaricus, but it is diagnosed most clearly by the 2 spines on the first peduncle segment of antenna 1 from the first two, and poorly setose antenna 2 and pereopods 5–7 for the latter, among many other characters. G. loeffleri might also show some similarities with G. crinicaudatus, G. komareki, G. birsteini, and G. rambouseki, but there are strong differences separating it from all these species as well. No much similar morphologies were seen among reported Russian and Central Asian freshwater Gammarus species to G. shirazinus and G. loeffleri. Among Chinese gammaridean fauna, G. qiani and G. translucidus shows the similarity of long setae on anterior pereopods 5–7 segments, still with shorter ones, but both of them have many other features differentiating the two species described here from Iran. SEM. The pioneer works on cuticular ultra-structure provided invaluable information for a list of gammarids including Gammarus pulex, G. agrarius, G. lordeganensis, G. bakhteyaricus, G. duebeni, G. lobifer, and G. balutchi (Khalaji-Pirbalouty & Sari 2006). Some of these species possess exclusively line (L) type, while others show scattered (S) type of pore arrangement on their cuticle surface polygons. In contrast, populations studied in the current investigations, including the two new species described here, as well as previously described G. zagrosensis and G. sepidannus (Zamanpoore et al. 2009) demonstrate both L- and Stype of pore arrangement on their cuticles. Additionally, SEM micro-photographs from the patterns of S-type pore distribution over the cuticle surface showed a special patchy or cluster pattern in G. shirazinus (Fig. 6C) in contrast to the random distribution in G. loeffleri (Fig. 6D). Although these two species showed overlapping ranges of pore line distances, the density of pores in both L-/ S-type polygons were higher in G. loeffleri cuticle surface (mean values: 108/ 176) than those of G. shirazinus (mean values: 64/ 131), indicating closer distances between pores as another factor of variety.



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Despite no ultra-structural data is available up to this time for the closest species compared earlier in this paper with G. shirazinus and G. loeffleri for further supporting of the macro-morphological evidences, existence of clearly different patterns, line and pore distances, and pore density among all these studied species sounds sufficient for deciding on a species-specific nature for their cuticular ultra-structure, leading to a better judgment of their taxonomic separation as good species. Comparisons of previously established/ recently described species and the two new species discussed here demonstrate additional sources of variation among Gammarus species, strengthening the suggestion of the usefulness of these ultra-structural features in gammaridean taxonomy, as well as in some other taxa (Elfimov, 1995; Grygier, 1995; Sari, 1997). Ecology. Effects of some aquatic environmental factors including temperature and salinity on the quality of life in various crustaceans have been extensively emphasized (Hagerman & Weber, 1981; Schmidt-Nielsen, 1997; Achituv & Cook, 1984; Spanopoulos-Hernándeza et al., 2005), most evidently as an increase in oxygen consumption and respiratory activity (Achituv & Cook, 1984; Allan et al., 2006; Chen & Nan, 1993; Isla & Perissinotto, 2004; Spanopoulos-Hernándeza et al., 2005). It has also been shown that macrozoobenthos community compositions can be significantly influenced by temperature (e.g. Živić et al., 2006). Our results showed that losing altitude at the south-eastern margins of Southern Zagros Region and the gradual merging of the Zagros Mountains with the hot arid plains in the east ends up in the formation of a novel type of habitats for amphipods – warmer springs and rivers – which are not potentially the favourite habitat of this group (Karaman & Pinkster, 1977). From the zoogeographical point of view, this could act as an environmental pressure, leading to the evolution of local adaptations, so that the settled marginal populations could withstand the new habitats. At the same time, they could act as a barrier, preventing central populations from expanding over the new warmer ranges. Hence it seems convenient enough to regard higher water temperatures combined in some occasions to salinity as an ecological barrier, as well as a physiological barrier, with respect to its secondary effect on solubility and higher demand of oxygen. Altogether, they could be strong enough for habitat separation, preventing inbreeding and gene flow among populations on two sides, thus promoting the processes of speciation as a result. This phenomenon has been shown in some species of Gammarus in which reproductive isolation was gained as a result of habitat specializations produced by differences in salinity which was secondarily fixed in the genes (Kolding, 1985; Rock et al., 2007). A same effect can be expected for temperature due to its considerable role on aquatic life. Regarded as polyphyletic, evolution of freshwater amphipods was an outcome of numerous habitat colonisations in continental waters (Väinölä et al., 2008). Divergent ecological changes could be capable of setting forceful effects on physiological and occasionally reproductive isolation of populations, fortifying altogether the formation of new species. Diversification and establishment of various endemic species of fresh water gammarids in the Southern Zagros Region described by Mateus and Mateus (1990), Stock et al. (1998), Zamanpoore et al. (2009), and the present work can be referred to as an example.

Acknowledgements We would like to thank D. Platvoet, Zoological Museum of Amsterdam, and P. Dworschak, Natural History Museum of Vienna, for loan material. Special thanks to W. Klepal and D. Gruber, University of Vienna for SEM facilities. Some of older samples were dedicated by K. Elmi, University of Shiraz. Grateful thanks go to A. Hatami for identifying aquatic macrophytes. H. Krail, Limnology lab, University of Vienna, and sampling group M. Qaed-Abdi, E. Nasr, and N. Yazdi, and chemistry technician Narges Ahmadi, FARSAGRES, are deeply acknowledged. Finally, we would like to dedicate our special appreciation to Dr. G. Karaman for his useful comments on the manuscript.

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