Two new sympatric Knipowitschia species (Telesotei: Gobiidae) from an eastern Mediterranean coastal lake - examples of different dispersal patterns?

Abstract

Two new species of the genus Knipowitschia, Knipowitschia byblisia sp. nov. and Knipowitschia caunosi sp. nov., are described from the coastal Lake Köycegiz, southwest Turkey. Knipowitschia byblisia sp.nov. is placed in a group of species with reduced cephalic lateral-line canals and reduced squamation. This species is characterized by the presence of axillary and caudal peduncular patches of scales, by a distinct reduced head canal system with only the postorbital section of the supraorbital canal developed and with longitudinal and transversal rows of free neuromasts in the interorbit. Knipowitschia caunosi sp. nov. is placed in a group of species with a fairly complete head canal system and with scales continuously extending along lateral midline from the axilla to the caudal peduncle. This species is characterized by long transverse rows of free neuromasts on the nape and on the cheek, by a characteristic pattern of the preorbital series of free neuromasts with the anteriormost row lacking and with a reduced number of transverse suborbital rows. Both species are isolated and their distributions restricted to a small brackish lake in the west of the Anatolian south coast. This record represents the southern and easternmost of the genus Knipowitschia in the Mediterranean region. The origin of these two species is discussed.

Full text

22 Accepted by M.T. Craig: 10 Oct. 2011; published: 30 Nov. 2011
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Copyright © 2011 · Magnolia Press
Zootaxa 3114: 22–30 (2011)
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Article
Two new sympatric Knipowitschia species (Teleostei: Gobiidae) from an eastern
Mediterranean coastal lake—examples of different dispersal patterns?
HARALD AHNELT
University of Vienna, Department of Theoretical Biology, Althanstrasse 14, 1090 Vienna, Austria. E-mail: harald.ahnelt@univie.ac.at
Abstract
Two new species of the genus Knipowitschia, Knipowitschia byblisia sp. nov. and Knipowitschia caunosi sp. nov., are
described from the coastal Lake Köycegiz, southwest Turkey. Knipowitschia byblisia sp.nov. is placed in a group of spe-
cies with reduced cephalic lateral-line canals and reduced squamation. This species is characterized by the presence of
axillary and caudal peduncular patches of scales, by a distinct reduced head canal system with only the postorbital section
of the supraorbital canal developed and with longitudinal and transversal rows of free neuromasts in the interorbit. Knip-
owitschia caunosi sp. nov. is placed in a group of species with a fairly complete head canal system and with scales con-
tinuously extending along lateral midline from the axilla to the caudal peduncle. This species is characterized by long
transverse rows of free neuromasts on the nape and on the cheek, by a characteristic pattern of the preorbital series of free
neuromasts with the anteriormost row lacking and with a reduced number of transverse suborbital rows. Both species are
isolated and their distributions restricted to a small brackish lake in the west of the Anatolian south coast. This record rep-
resents the southern and easternmost of the genus Knipowitschia in the Mediterranean region. The origin of these two spe-
cies is discussed.
Key words: Gobiidae, Knipowitschia, new species, brackish lake, dispersal, Mediterranean region, Turkey
Introduction
Three species of the genus Knipowitschia Iljin are known from continental Aegean Anatolia, the euryhaline Knip-
owitschia caucasica (Berg) and the two freshwater species K. ephesi Ahnelt and K. mermere Ahnelt (Ahnelt 1995;
Van Neer et al. 1999; Miller 2004). Contrary to K. caucasica the other two species are endemic and only known
from isolated freshwater habitats (Ahnelt 1995; Ahnelt et al. 1995; Kovacic 2005).
The River Degrimen was believed to be the southernmost limit of the distribution of Knipowitschia species in
western Anatolia (Ahnelt 1995; Bianco et al. 1996). But a sample of gobies from Lake Köycegiz deposited in the
Ichthyological Collection of the Zoologisches Museum für Naturkunde der Universität Hamburg document for the
first time the occurrence of the genus Knipowitschia in the transition from the Aegean region to the eastern Medi-
terranean. These specimens represent the southern and easternmost record of Knipowitschia in the Mediterranean
region and the first on the southern coast of Anatolia.
Material and methods
Counts and measurements follow Miller (1988). Measurements were taken with the aid of an electronic calliper.
The terminology of the cephalic lateral-line canals follows Ahnelt (2001), those of the canal pores Akihito (1986).
Free neuromasts are listed by innervation categories for Gobiidae (Ahnelt & Bohacek 2004). The terminology of
the free neuromasts of the lateral-line system follows Sanzo (1911) except for the transversal oculoscapular row tr
(Ahnelt 2001) and the transversal interorbital row w (Ahnelt et al. 2000). Because the topography of free neuro-
masts generally resembles that of Knipowitschia caucasica (Miller 2004) only the characteristic differences are
described. Characters of the holotype are indicated by asterisks in counts and by brackets in measurements. Sex

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TWO NEW SPECIES OF KNIPOWITSCHIA FROM TURKEY
was determined by the shape of the urogenital papilla, uniformly wide in females, narrowing at the tip in males.
The types are deposited in the Ichthyological Collection of the Zoologisches Museum für Naturkunde Hamburg
(ZMH), Germany.
Knipowitschia byblisia sp. nov.
Byblis Goby
Figures 1–2
Synonymy. Pomatoschistus (Bubyr) caucasicus kosswigi. Ladiges 1964: 212 (part).
Holotype. ZMH 2175:1, female, 21.1 mm SL; Turkey, Lake Köycegiz, 36°55’N, 28°40’E; collected 31. October
1946.
Paratypes. ZMH 2175:2–2175:6, five females, 18.5–25.4 mm SL; ZMH 2175:7–2175:8, four males, 21.5–
25.2 mm SL; ZMH 2175:9–2175:13, five juveniles, 13.4–14.6 mm SL; same as holotype.
Etymology. Named for the mythological figure Byblis, twin sister of Caunos who founded the ancient city
Caunos at the Anatolian southwest coast. The ruins of this city are located close to Lake Köycegiz.
Diagnosis. A Knipowitschia with (1) scales in two patches in the axillary area and on the caudal region respec-
tively; (2) rear part of the caudal peduncle, immediately anterior to the caudal fin, naked; (3) head canals reduced to
two short postorbital canals; (4) three rows of free neuromasts in the interorbit, a paired longitudinal row and a
short transverse row; (5) internal preoperculo-mandibular row extending postero-dorsally replacing the preopercu-
lar canal, at least in part of the population; (6) male colouration with 8–9 dark vertical bars on the sides, dark blotch
on the rear of the first dorsal fin, and (7) first dorsal fin with six to seven rays.
FIGURE 1. Top: Knipowitschia caunosi, ZMH 2175:16, holotype, female, 26 mm SL. Bottom: Knipowitschia byblisia, ZMH
2175: 1, holotype, female, 21.1 mm SL. Both from Lake Köycegiz, Turkey.
Description. Measurements of the holotype (21.1 mm SL) and nine adult paratypes (18.5–25.4 mm SL) as per-
cent of standard length (range and mean in parentheses, values of holotype in brackets): head length 26.1–30.4
(27.8) [30.8], head width 12.2–12.6 (12.3) [11.4], distance from snout to origin of first dorsal fin 37.4–42.1 (39.5)
[43.1], distance from snout to origin of second dorsal fin 56.7–60.1 (58.9) [57.9], distance from snout to anus 53.2–
56.4 (54.9) [53.7], distance from snout to origin of anal fin 57.4–59.3 (58.8) [58.1], distance from snout to origin of
pelvic fin 30.8–33.1 (31.6) [33.2], caudal peduncle length 22.3–26.1 (25.3) [23.2], length of first dorsal fin base
9.8–10.6 (10.3) [10.9], length of second dorsal fin base 15.6–17.5 (16.8) [17.1], length of anal fin base 13.3–15.0

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24 · Zootaxa 3114 © 2011 Magnolia Press
(14.1) [15.2], caudal fin length 21.9–23.2 (22.4) [22.1], pectoral fin length 21.3–23.6 (22.8) [21.3], pelvic fin
length 20.4–22.9 (22.0) [20.6], body depth at pelvic fin origin 18.6–19.7 (19.1) [18.5], body depth at anal fin origin
13.3–15.0 (14.2) [14.2], body width at anal fin origin 8.3–10.2 (9.4) [9.1], caudal peduncle depth 10.2–11.6 (10.7)
[9.6] distance from origin of pelvic fin to anus 25.5–28.1 (26.8) [24.9]; as percent in caudal peduncle length: caudal
peduncle depth 37.6–43.4 (41.8) [44.7]; as percentage of head length: snout length 20.6–23.5 (21.8) [20.0], postor-
bital length 46.1–50.0 (48.4) [49.2], eye diameter 25.5–28.6 (26.7) [26.2], cheek depth 17.5–19.0 (18.1) [17.9]; as
percentage of eye diameter: interorbital width 55.5–62.2 (58.9) [54.7]; as percentage of distance from origin of pel-
vic fin to anus: pelvic fin length 73.6–90.6 (92.7) [99.0].
Counts (values of the holotype are indicated by *): first dorsal fin VI–VII (VI*: 14, VII: 1); second dorsal fin I/
7–9 (7: 1, 8*: 12, 9: 2); anal fin I/7–8 (7*: 9, 8: 6); pectoral fin 15–17 (15*: 6, 16: 8, 17: 1); pelvic fins (united,
forming disc) I/5+5/I, fifth ray longest; caudal fin 15 segmented and 13 branched rays.
The squamation is reduced to two patches of ctenoid scales, one in the axillary area, the second on the caudal
peduncle (n = 7*). In two other specimens these patches are connected by a narrow band of scales in lateral mid-
line. Specimens <15 mm SL are not considered. The scales of the posterior patch do not extend to the rear end of
the caudal peduncle, leaving a narrow naked area immediately anterior of the caudal fin.
Lateral-line system (Figure 2): Head canals distinctly reduced. Posterior oculoscapular and preopercular canals
absent. Only remnants of the anterior oculoscapular canal are left, i.e. the postorbital sections of the supraorbital
canal with the pores D and F present. Both canal remnants are not fused in the midline.
FIGURE 2. Neuromast patterns and canal pores on the head of Knipwitschia byblisia, ZMH 2175:1, holotype, female, 21.1
mm SL. PN, posterior nostril; for other terminology see text. Scale bar = 1 mm.
Rows and, in parentheses mean of numbers of free neuromasts (sensory papillae) from six type specimens with
conspicuous head neuromasts which are important to separate this species from K. caunosi and from other
congeners.
Innervation by the anterior lateral-line nerve: (1) supraorbital: longitudinal rostral row s (11.5) a continuous
row ending anteriorly close to anterior nostril; longitudinal interorbital row p (6.2) replacing interorbital section of
the supraorbital canal; transversal row w (5.8), continuous (n = 4) or divided in a left and a right section (n = 2). (2)
Infraorbital: longitudinal row a (8.5) with one to two transverse rows (1.8); transversal row s3 (2.9) internal to

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TWO NEW SPECIES OF KNIPOWITSCHIA FROM TURKEY
supraorbital row s and close to upper lip. (3) Hyomandibular longitudinal row b (9.8) anteriorly not reaching below
orbit; longitudinal row i (37.8) at least with two (n = 4) or with 3–5 (n = 2) neuromasts in course of the reduced pre-
opercular canal. (4) Otic transversal row tra (3.7) short and confluent with supraorbital row n. Innervation by the
posterior lateral-line nerve: (1) supratemporal transversal row tr (6.7) short, not reaching close to longitudinal row
m (4.1). (2) Posterior dorsal row y (1.0) present. Posterior lateral transverse axillary row as1 (12.1) long, dorsally
extending well above the longitudinal axillary rows la.
The possession of a transverse row of free neuromasts in the interorbit was believed to be a feature to separate
Hyrcanogobius Iljin 1928 from Knipowitschia (Miller 2004).
Coloration in preserved specimens. Description of colouration is based on preserved material. All specimens
are pale fawn. Markings on the body are very indistinct, possibly due to preservation. Two males with about eight
to nine faded stripes and blotches along lateral midline and a distinct dark spot in the rear of the first dorsal fin
restricted to the area between fifth and sixth fin ray, not extending to the end of the fin membrane. Indistinct dark
blotches in lateral midline of females but very faded. No dark blotch on chin in females.
Knipowitschia caunosi sp. nov.
Caunos Goby
Figures 1, 3
Synonymy. Pomatoschistus (Bubyr) caucasicus kosswigi. Ladiges 1964: 212 (part).
Holotype. ZMH 2175:16, female, 26.0 mm SL; Turkey, Lake Köycegiz, 36°55’N, 28°40’E; collected 31. October
1946.
Paratypes. ZMH 2175:17, one male, 25.6 mm SL, ZMH 2175:18, one female, 26.6 mm SL; same as holotype.
Etymology. Named for the mythological figure Caunos, twin brother of Byblis and founder of the ancient Car-
ian city Caunos. The ruins of this city are located close to Lake Köycegiz.
Diagnosis. A Knipowitschia with (1) scales from the axilla in midline to the caudal peduncle, extending onto
base of caudal fin; (2) anterior oculoscapular canal present, ending anteriorly with a pair of interorbital pores; (3)
preopercular canal present with two terminal pores; (4) external preorbital neuromast row extending anteriorly
close to the upper lip; (5) anterior preorbital row absent; (6) anterior transverse row long, extending on the nape;
(7) pelvic frenum with smooth free edge; (8) male colouration without vertical bars or striae, dark blotch in the rear
of the first dorsal fin; first dorsal fin rays V–VI.
Description. Measurements of the holotype (25.9 mm SL) and two paratypes (25.6–26.5 mm SL) as percent of
standard length (range and mean in parentheses, values of holotype in brackets): head length 25.7–25.8 (25.8)
[25.5], head width 11.3–12.5 (11.9) [11.9], distance from snout to origin of first dorsal fin 36.2–37.5 (36.9) [37.6],
distance from snout to origin of second dorsal fin 54.7–55.8 (55.3) [55.2], distance from snout to anus 50.8–55.5
(53.2) [54.4], distance from snout to origin of anal fin 55.5–59.2 (57.4) [58.3], distance from snout to origin of pel-
vic fin 29.3-31-3 (30.3) [29.7], caudal peduncle length 28.3–29.7 (29.0) [29.7], length of first dorsal fin base
8.7+10.7 (9.7) [8.9], length of second dorsal fin base 14.7–16.8 (15.8) [15.1], length of anal fin base 12.9–14.4
(13.7) [12.4], caudal fin length 20.4–22.3 (21.4) [damaged], pectoral fin length 20.0–22.3 (21.2) [20.1], pelvic fin
length 22.3+23.0 (22.7) [22.0], body depth at pelvic fin origin 19.6–19.9 (19.8) [19.3], body depth at anal fin origin
15.5–16.8 (16.2) [16.2], body width at anal fin origin 9.8–10.2 (10.0) [10.0], caudal peduncle depth 9.0–9.8 (9.4)
[9.3], distance from origin of pelvic fin to anus 23.8–26.4 (25.1) [28.0]; as percent in caudal peduncle length: cau-
dal peduncle depth 32.0–32.9 (32.5) [31.2]; as percent of head length: snout length 24.2–26.5 (25.4) [22.9], postor-
bital length 46.6–55.1 (50.9) [51.5], eye diameter 24.2–27.9 (26.1) [22.7], cheek depth 20.6–22.7 (21.7) [17.7]; as
percent of eye diameter: interorbital width 47.4–50.0 (48.7) [53.3]; as percent of distance from origin of pelvic fin
to anus: pelvic fin length 85.7–95.2 (90.4) [82.9].
Counts (values of the holotype are indicated by *): first dorsal fin V–VI (V: 2, VI*: 1); second dorsal fin D2 I/
8; A I/7–8 (7: 1, 8*: 29); pectoral fin 15–17 (15*: 2, 17: 1); pelvic fin complete, pelvic fins (united, forming disc) I/
5+5/I, fifth ray longest; caudal fin 15 segmented and 13 branched rays; scales in lateral series 30 –32 (30: 1, 32*:
2), scales in transversal series 8.
The body squamation is typical for a Knipowitschia species with head, nape, back to origin of second dorsal fin
and breast naked. The ctenoid scales extend posteriorly from the base of the pectoral fin on the base of the caudal fin.

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Lateral-line system (Fig. 3): The anterior oculoscapular canals are fused in the midline at a single pore D with
paired interorbital pores C and postorbital pores F and H or not fused in midline with two pores in position of pore
D and with short remnants of the interorbital canals anterior to pore D, as a short canal or as furrow. A preopercular
canal is developed, with pores M and O, in one specimen or just as an open furrow in the other two.
FIGURE 3. Neuromast patterns and canal pores of Knipwitschia caunosi, ZMH 2175:16, holotype, female, 26 mm SL. PN,
posterior nostril; for other terminology see text. Scale bar = 1 mm.
Rows and, in parentheses, mean of numbers of free neuromasts (sensory papillae) from the three type speci-
mens which are important to separate this species from K. byblisia and from other congeners.
Innervation by the antertior lateral-line nerve: (1) supraorbital: longitudinal rostral row s (10.3) a continuous
row ending anteriorly close to upper lip; longitudinal interorbital row p absent, also in one specimen where the
interorbital section of the supraorbital canal is not closed but present as a deep furrow. (2) Infraorbital: longitudinal
row a (8.3) with two to three short transverse rows; transversal row s3 absent. (3) Hyomandibular longitudinal row
b (11.7) anteriorly reaching below orbit; longitudinal row i (32.3) ending close to pore O of the preopercular canal.
(4) Otic transversal row tra (6.6) confluent with supraorbital row n and ventrally extending two third of distance to
hyomandibular longitudinal row b.
Innervation by the posterior lateral-line nerve: (1) supratemporal transversal row tr (13.5) long, reaching close
to longitudinal row m (5.3). (2) Posterior dorsal row y absent. Posterior lateral transverse axillary row as1 (8) short,
dorsally not extending above the longitudinal axillary rows la.
Coloration in preserved specimens. Description of colouration is based on preserved material. All specimens
are pale fawn. No markings on the body have been identified. The first dorsal fin in males has a distinct dark spot
in the rear, extending from fifth ray to the end of the fin membrane. No dark blotch on chin in females.
Remarks. Distribution: Six species of the Ponto-Caspian genus Knipowitschia, K. caucasica, K. ephesi, K.
mermere, K. thessala (Vinciguerra), K. byblisia sp. nov. and K. caunosi sp. nov. occur in peripheral habitats around
the Aegean Sea (Ahnelt et al. 1995; Miller 2004; this study) (Figure 4). These species are primary-like freshwater
fishes that evolved from an ancient euryhaline ancestor of Paratethys and show a similar pattern of distribution to
that of primary freshwater fishes (Bianco et al. 1996). They likely dispersed the same way, were affected by con-
nection events during the last glacial maxima, and may have found refuges in suitable coastal habitats after sea
level rises (Ahnelt 1995; Bianco et al. 1996; Durand et al. 2003; Reyjol et al. 2007).
Knipowitschia byblisia and K. cauonsi, only known from Lake Köycegiz, represent the southern and eastern-
most autochthonous record of this genus in the Mediterranean region and the first record of Knipowitschia in the
transition of the Aegean to the eastern Mediterranean. Note that K. caucasica has been introduced unwittingly to
Anatolian lakes (Van Neer et al.
1999). Recently, K. caucasica has also been reported from Lake Köycegiz (Balik
et al. 2005). Possibly this species has been misidentified (see Ladiges 1964), or K. caucasica has been introduced

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with fry of Cyprinus carpio Linnaeus, as has been previously described for Anatolian lakes (Van Neer et al. 1999).
All carp fry stocked before 1995 came from fish farms at the River Evros/Meric (Thracia), where K. caucasica
occurs naturally (Van Neer et al. 1999). Furthermore, a land-based fish farm, a possible source of introduction, is
located close to the Köycegiz tributary Yuvarlakcay (Taseli 2009). Stocking of C. carpio for recreational fishing
started in the 1950s/1960s (Innal & Erk’akan 2006). Nevertheless, the material under investigation was collected in
1946, which confirms the autochthonous origin of K. byblisia and K. caunosi in Lake Köycegiz.
FIGURE 4. Geographical distribution of the genus Knipowitschia in the Aegean Region. Open circle, K. thessala; filled circle,
K. caucasica, open triangle, K. mermere; filled triangle, K. ephesi; star, Lake Köycegiz, type locality of K. byblisia and K.
caunosi.
Ecology. No ecological data are associated with the sample of Knipowitschia from Lake Köycegiz and the
exact collection site is unknown. This small lake is characterised by brackish water (Akin et al. 2005), which
would have favoured a euryhaline stock of Knipowitschia. Lake Köycegiz, a former bay of the Mediterranean Sea,
was already in existence 4,000 BP. About 2,000 BP the harbour of the ancient city of Kaunos, located close to the
lake, lost its connection to the open sea (Brückner 1997).
Origin. The occurrence of K. byblisia and K. caunosi in the transition area from the Aegean region to the east-
ern Mediterranean can be explained by two major events: the Messinian salinity crisis of the Mediterranean Sea at
the end of the Miocene (about 5.6 million years ago) and the Pleistocene glaciations (between 650,000–11,000
years ago). During the latter event, the salinity of the Black Sea varied substantially (Yanko-Hombach et al. 2007).
At the end of the glacial maxima outflow of low salinity (2–6 ppt) or even fresh water connected the Black Sea

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with the Mediterranean Sea (summarized in Yanko-Hombach et al. 2007). This enabled immigration of the autoch-
thonous Ponto-Caspian ‘sand goby’ genus Knipowitschia, in particular the euryhaline K. caucasica sensu Miller
(2004) to the Aegean region where this species inhabits circum Aegean lagoons and freshwater habitats (Econo-
midis & Miller 1990; Ahnelt et al. 1995, Bianco et al. 1996). Gene flow between these populations is limited or
prevented by marine conditions (33 ppt) and by the counter-clockwise surface currents in the Aegean Sea (Hamad
et al. 2005). A similar distribution in isolated freshwater habitats in the Mediterranean basins is known from Gas-
terosteus aculeatus Linnaeus. These populations apparently trace back to a stock of G. aculeatus which inhabited
the Mediterranean Sea during Pleistocene glacial periods (Paepke 2002).
The second event, the salinity crisis of the Mediterranean, is characterized by a drastic drop in sea level and
subsequently the establishment of brackish conditions in the “Lago Mare” phase of the Mediterranean basins
(Penzo et al. 1998; Huyse et al. 2004). The brackish fauna included many species of Paratethys origin (Popov et al.
2006) reaching the Mediterranean basins by spill-over (Rögl 1998). These fresh and brackish environments
enabled the dispersal of euryhaline and freshwater fishes (Reyjol et al. 2007). After the formation of the modern
Mediterranean Sea about 5.3 million years ago only isolated populations of a euryhaline ancestor of the Mediterra-
nean Knipowitschia populations survived in restricted areas such as estuaries and brackish lagoons (Penzo et al.
1998; Huyse et al. 2004). Possibly such an ancestor gave rise to the Adriatic and the Aegean Knipowitschia stocks,
and isolation in estuaries, brackish lagoons and peripheral freshwaters clearly favoured speciation. Similar relicts
of the Mediterranean ‘Lago Mare’ fish fauna are the recent Aphanius and Salaria species (Perdices et al. 2000;
Hrbek & Meyer 2003).
Knipowitschia byblisia and K. caunosi occur sympatrically in a small coastal lake. Such a sympatric occur-
rence of two closely related Knipowitschia species is rare (Miller 2004), but is possibly based on different radiation
scenarios. K. byblisia, the most specialized and in its paedomorphic features similar to K. ephesi and K. mermere
from eastern Aegean freshwater habitats (see below), could trace back to an ancient Knipowitschia stock from the
Mediterranean ‘Lago Mare’ phase. K. caunosi, more similar to K. caucasica, apperently radiated from a K. cauca-
sica stock reaching the area of the Köycegiz lake during melting phases of Pleistocene glaciations. Possibly these
different radiation scenarios also explain the paraphyletic position of the genus Knipowitschia proposed by Penzo
et al. (1998) and Hyuse et al. (2004).
Morphology. Knipowitschia caunosi, and to a lesser extent K. byblisia, resemble the euryhaline eastern
Aegean K. caucasica stock but differ in squamation and in the type of reduction of the head canals. Differences are
distinct, particularly with K. caucasica, a species of larger body size and with completely developed head canals of
the lateral line system, with fusion of the anterior oculoscapular canals in midline and posterior oculoscapular canal
present (Miller 2004). With reduction of body size, in head lateral-line canals and squamation,
K. byblisia shows
similar paedomorphic features to K. ephesi and K. mermere (Ahnelt 1995; Ahnelt et al. 1995) or the freshwater
species from the Adriatic region (Kovacic 2005; Kovacic & Sanda 2007). All these species are miniature as adults
(< 28 mm SL) and characterised by high morphological uniformity due to paedomorphosis. Small body size offers
increased habitat opportunities, a fine subdivision of the environment and it enables survival in small habitats with
limited or fluctuating resources (Dial & Marzluff 1988; Miller 1996). Additionally, small body size is often linked
to short generation time (Miller 1996), which favours diversity and speciation (Martin & Palumbi 1993).
Completion of the life cycles of small gobiid fishes, in isolated and small habitats seems to favour reduction or
loss of squamation and of the head lateral-line canals (Ahnelt 1995; Ahnelt et al. 2004; Kovacic & Sanda 2007). If
replaced by free neuromasts, the loss of head canals may also contribute to increased sensitivity of the lateral line
system as discussed for Eucyclogobius newberryi (Girard) (Ahnelt et al. 2004; Earl et al. 2010). This small north-
east Pacific gobiid shows similar reductions of the supraorbital canal to K. byblisia (Ahnelt et al. 2004, Fig. 3).
Clearly similar habitat and life history, combined with low dispersal, favour similar morphological characteristics,
such as small body size, reduced squamation and reduced head lateral-line canals. Such scenarios apparently
occurred independently in E. newberryi and in the Adriatic and Aegean clades of Knipowitschia. It is therefore
likely that such a suite of characteristics is a general indicator of gobiid species adapted to small habitats and there-
fore of special interest for evolutionary studies and for conservation.
Conservation. The fresh water fish fauna of continental western Anatolia is critically endangered in coastal
areas by anthropogenic effects (Ahnelt et al. 1995; Innal & Erk’akan 2006). Agriculture, aquaculture, hydro power
stations and tourism greatly affect the habitats of fresh water fishes. The native fish fauna of Lake Köycegiz is
affected by pollution and increasing tourism, as well as by the introduction of non-native fish species (Innal &

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TWO NEW SPECIES OF KNIPOWITSCHIA FROM TURKEY
Erk’akan 2006, Taseli 2009). There is also a single record of K. caucasica in Lake Köycegiz (Balik et al. 2005). If
this is correct, this congener, with similar ecological demands and larger body size, could out-compete the two
native species. Additionally, introducing alien species, e.g. Tilapia zillii (Gervais 1848) (Akin et al. 2005; Balik et
al. 2005) will increase predation pressure. Known only from Lake Köycegiz, it is likely that K. byblisia and K.
caunosi are threatened by competition and predation pressure imposed by introduced species.
Acknowledgements
For the loan of the material and for his hospitality I thank R. Thiel, head of the Ichthyological Collection of the
Zoologisches Museum für Naturkunde der Universität Hamburg (ZMH). Special thanks go to M. Petrasko for his
technical support and to P. Humphries for improving the English.
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