Palaeozoology Research Article ACTA ZOOLOGICA BULGARICA Acta zool. bulg., 67 (2), 2015: 283-290 Porzana botunensis sp. n., a New Early Pleistocene Crake (Aves: Rallidae) from Bulgaria Zlatozar Boev National Museum of Natural History, Bulgarian Academy of Sciences, 1 Tsar Osvoboditel Blvd., 1000 Sofia, Bulgaria; E-mail: boev@nmnhs.com; zlatozarboev@gmail.com Abstract: A new species of crake of the genus Porzana is described from the site of Varshets (MN 17). Porzana botunensis sp. n. is the second rallid species described from the site. The holotype is a distal half of left tarsometatarsus. The species may be diagnosed using the following characters: an Early Pleistocene crake, differing from the recent P. porzana in the: (1) more distally positioned inception of trochlea metatarsi II; (2) shallower edges of the condyles of the trochlea metatarsi II; (3) clearly developed edge (linea) between the facies dorsalis and facies subcutanea lateralis; (4) deeper facies lateralis caudalis and (5) relatively wider incisura intertrochlearis lateralis. Keywords: Early Pleistocene, Villafranchian fauna, fossil vertebrates, Tertiary birds, family Rallidae, Gruiformes, Balkan Peninsula, Bulgaria Introduction The Early Pleistocene (Villanyian, MN 17 zone) vertebrate fauna of the studied site and its taphonomy have been characterised in Boev (2010). This unique ponor provided a large number of avian fossils of over 74 taxa. Studies at this site started in 1988. It is the type locality of a series of species. The character of the paleoenvironment, according to discovered paleoavifauna, was determined as forest-steppe (Boev 1995; 1999). The Pra-Botunya River passed near the site, while at present it is about 1 km away eastwards. The aim of the present article is to describe a new Early Pleistocene crake of the genus Porzana Vieillot, 1816 recorded the site of Varshets (MN 17). Material and Methods The material was collected through screening and washing of sediments. It was identified using reference to comparative bird collections of the ISEA, NHM, NMNHS and UCBL (see abbreviations below). Characteristic measurements of distal tarsometatarsus in Rallidae: a – width of the diaphysis in the middle; b – minimal thickness of the diaphysis; c – width of tr. m. III; d – diameter of tr. m. III. All measurements are given in mm. Total length of the fragment: 15.3 mm. Abbreviations Anatomical: f. m. – fossa metatarsi I; f. l. c. – fovea ligamenti collateralis; tr. m. – trochlea metatarsi; Institutional: ISEA – Institute of Systematics and Evolution of Animals (Krakow); NHM – Natural History Museum, formerly British Museum (Natural History, Tring); NMNHS – National Museum of Natural History (Bulgarian Academy of Sciences, Sofia); UCBL – University Claude Bernard, Lyon 1 (Lyon). Taxonomy Gruiformes Bonaparte, 1854 Rallidae Vigors, 1825 Porzana Vieillot, 1816 Porzana botunensis sp. n. Holotype: No NMNHS 301 distal half of a left tarsometatarsus of excellent preservation (Fig. 1), collections of the Fossil and Recent Birds Department of the National Museum of Natural History, Bulgarian Academy of Sciences, No NMNHS 301. Collected on 5 June 1992 by Zlatozar Boev. No additional material was collected. Locality: A ponor in a rocky hill, 6 km NE of Varshets (43.13 N, 23.17 E). Stratigraphic position: Unconsolidated, unstratified sediments accumulated in the filling of clay 283 Boev Z. Fig. 1. Porzana botunensis sp. n. – tarsometatarsus dex. ad., holotype, Early Pleistocene, Varshets (Bulgaria), No NMNHS 301 (left) and Porzana porzana ♂ ad., recent, Sofia Region (Bulgaria) No NMNHS 1/1989 [889] (right): cranial view – A; caudal view – B; medial view – C; lateral view – D; ventral view – E. Scale bar = 1 cm. (Photograph: Assen Ignatov, NMNHS) terra rossa. The fossil bones are broken, sometimes making a kind of bone breccia. Chronostratigraphy: Middle Villafranchian. The associated fauna of large mammals attributes the site to the MN 17 zone (SpaSSov 1997) according to the chronostratigraphical system of Mein (1990). popov (2001) determines the time span of Varshets between 2.04 Ma (Stranzendorf I) and 2.4 Ma (Stranzendorf D). Etymology: The name “botunensis” is given after the name of the Botunya River, the main geologic agent for the formation of the karstic landscape on the terrain of the locality. 284 Diagnosis: An Early Pleistocene crake, differing from the recent P. porzana in the: (1) more distally positioned inception of trochlea metatarsi II; (2) shallower edges of the condyles of the trochlea metatarsi II; (3) clearly developed edge (linea) between the facies dorsalis and facies subcutanea lateralis; (4) deeper fovea ligamenti collateralis; and (5) relatively wider incisura intertrochlearis lateralis. For measurements and comparative material examined see Table 1. The total length of the fragment is 15.4 mm. The finding shows all features characteristic for the tarsometatarsal bone of the small-sized taxa of Porzana botunensis sp. n., a New Early Pleistocene Crake (Aves: Rallidae) from Bulgaria the family Rallidae, including almost parallel condyles of the trochlea metatarsi II, III and IV, and much higher position of the trochlea metatarsi II. Comparison with some recent taxa of other genera of the family Rallidae The specimen from Varshets differs from: Rallus aquaticus Linnaeus, 1758 in the smaller size, more asymmetrical tr. m. III in cranial view, shallower relief of the tr. m. IV and tr. m. III, and flat cranial surface (facies dorsalis) of the distal half of diaphysis (Fig. 2); Laterallus leucopyrrhus (Vieillot, 1819) in the correlation between the diameters of tr. m. III and tr. m. IV; Sarothrura elegans (A. Smith, 1839) in the thicker and right shaft of diaphysis, bigger diameter of tr. m. II, deeper with clear edges f. m. I, and less developed relief of the tr. m. II; Amaurornis flavirostris (Swainson, 1837) in the smaller size, less developed relief on tr. m. IV, and deeper and better shaped fossa of foramen vasculare distale; Crex crex (Linnaeus, 1758) in the smaller size (Table 1), considerably more protruding trochlea metatarsi III beyond the trochlea metatarsi IV in lateral view, and parallel condyles of tr. m. II compared with tr. m. III and tr. m. IV; Crex egregia (W. K. H. Peters, 1854) in the smaller size, more graduate instead of sharp, transition in medial view of the profile from tr. m. II to the diaphysis; Canirallus kioloides (Pucheran, 1845) in the relatively narrower tr. m. III (measurements c : d); Gallinula melanops (Vieillot, 1819) in the smaller size, lower relief of tr. m. II, and sharper, but not round, profile of tr. m. II in medial view; Porphyrio alleni Thomson, 1842 in the round, but not elongated foramen vasculare distale, smaller size and flater, but not bow-like cranial surface in the middle of the diaphysis; Porphyrio martinica (Linnaeus, 1766) in the smaller size, because that species is very close dimensionally to P. alleni. According to metrical data by JánoSSy (1991) the specimen from Varshets differs from Rallicrex polgardiensis Jánossy, 1991 not only osteometrically, but also morphologically. He writes: “The measurements (i.e. of R. polgardiensis) are absolutely larger than the same ones of the smaller rallid of Polgardi (Porzana estramosi veterior Jánossy, 1991).” (p. 22). The tarsometatarsus of R. polgardiensis stands closely to the one of Crex crex and shows conspicuous robustness (JánoSSy, 1991). The Upper Oligocene or Lower Miocene (Aquitanian or Tortonian) Paraortygometra porzanoides (Milne-Edwards, 1869), described from France, besides its close dimensional and morphometrical affinity to the genus Porzana (Olson, 1977), could be excluded from our comparison because of the considerable chronostratigraphical differences: MN 4b in Dolnice (Czech Republic; MlíkovSký, 1996 b), MN 2a in Saint-Gerand-Le-Pyu and MN 7-8 in La Grive-Saint-Alban (France; Cheneval, 1996). As de Pietri and Mayr (2014) have found, Paraortygometra is a stem-representative of Rallidae, another reason to be excluded from the comparisons. We exclude the similarly-sized species bandbellied crake (Porzana paykullii Ljungh, 1813), a species with East-Asian distribution (both breeding and wintering ranges). The genus Porzana (P. porzana was compared) has more laterally protruded lateral edge of the distal diaphysis of tmt. above the epiphysis in the area of tr. m. IV in cranial view than in Zapornia (Z. pusilla and Z. parva were compared). In Z. pusilla and Z. parva it is straighter instead of curved outwards. In this respect the fossil specimen resembles Porzana. The flat cranial surface of diaphysis within the 3rd fourth of its length, round shape of foramen vasculare distale and its position on the cranial surface on the distal epiphyisis also refer NMNHS 301 to genus Porzana. On the other hand, the specimen NMNHS 301 has a very clearly developed edge (linea) between Fig. 2. Ventral view of distal left tarsometatarsus in some West-Palaearctic rallids (left to right): Porzana botunensis sp. n., Z. parva, Rallus aquaticus, Crex crex, Z. pusilla (Drawing: Vera Hristova) 285 Boev Z. facies dorsalis and facies subcutanea lateralis in contrast to all compared recent species of Porzana. Recent and fossil crakes of Porzana and Zapornia Summarising data on the fossil record of Rallidae, olSon (1985) states that rails occur regularly in the Neogene deposits of the Northern hemisphere. CraCraft (1973) and olSon (1977) list no data on fossil taxa in the genus Porzana. A number of endemic fossil/subfossil late Quaternary (Holocene) extinct crakes of genus Porzana have been described also from the Pacific tropical islands: Porzana rua Steadman 1986 – Holocene of Mangaia, Cook Islands, Pacific Ocean; Porzana ziegleri Olson & James 1991 – Holocene of Oahu, Hawaiian Islands; Porzana menehune Olson & James 1991 – Holocene of Molokai, Hawaiian Islands; Porzana keplerorum Olson & James 1991 – Holocene of Maui, E Hawaiian Islands; Porzana ralphorum Olson & James 1991 – Holocene of Oahu, Hawaiian Islands; Porzana severnsi Olson & James 1991 – Holocene of Maui, E Hawaiian Islands; Zapornia astrictocarpus (Olson 1973) – Late Holocene (extinction after 1502 AD) of Str. Helena, S Atlantic Ocean. Most of them are flightless forms of very restricted island ranges. According to Bocheński (1997) 12 Neogene and one Quaternary rallids have been described from Europe, but only one (Porzana estramosi Jánossy, 1979) is referred to the genus Porzana. According MlíkovSký (2002) the genus Porzana is known from Europe since the Middle Miocene (MN 6-8) from Hungary. The second oldest record came from Varshets (present find listed as Porzana sp., but erroneously dated MN 18, instead of MN 17). Mlíkovský (2002) also states that P. estramosi is the only fossil species of the genus in Europe. Comparison with fossil species of the genus Porzana The fossil record of the genus spans over the period from the Middle Miocene MN 6-8 to Late Pleistocene (MlíkovSký 2002). Brodkorb (1967) lists three fossil species of Porzana: P. lacustris Brodkorb, 1958 (Lower Pleistocene, Idaho, USA); P. auffenbergi Brodkorb, 1954 (Middle Pleistocene, Florida, USA); and P. guti (Brodkorb, 1952) (Middle Pleistocene, Florida, USA). Porzana auffenbergi is now placed in Rallus. In addition, tree other fossil taxa are recognised: P. estramosi Jánossy, 1979 (Middle? Miocene – Early Pliocene of Hungary); P. risilla (Kurochkin, 1980) (Middle Miocene or Middle Pliocene of Mongolia); P. piercei Olson, Wingate, 2000 (Late Pleistocene of Bermuda, W Atlantic). Porzana risilla, as Kurochkin 286 (1985) stated is similar in size to P. pusilla and could be excluded from the comparisons. Metrically the find from Bulgaria lies in the range of genus Porzana. Porzana estramosi from the Middle Pliocene (“Postpannonian Pliocene”) of Northern Hungary was described by a distal fragment of left tarsometatarsus (JánoSSy, 1979). It is a species “smaller than any hitherto described fossil and recent continental forms” (p. 20). As it is seen from Table 1, the specimen from Bulgaria has intermediary for the genus Porzana measurements and may not be referred dimensionally to P. estramosi. The photograph 4-8 on p. 18 of the paper by JánoSSy (1979) shows that both medial and lateral edges of the base of diaphysis above the tr. m. III in P. estramosi form a clear narrowing, that is not present in P. botunensis sp. n.. Later D. Jánossy describes a Late Miocene subspecies of P. estramosi: P. e. veterior Jánossy, 1991, based chiefly on its smaller dimensions. He establishes that P. e. veterior has longer and slenderer tarsometatarsi in comparison with Z. parva: a character that also excludes the taxonomic identity of the Bulgarian specimen and P. estramosi. MlíkovSký’S (1996c) summary data show that no fossil record of genus Porzana is known from Europe except from the sites of Varshets (Bulgaria), Polgardi and Osztramos 9 (Hungary). tyrBerg (1998) cites the opinion of J. Mlíkovský who states that the Middle Pleistocene (MNQ 21) finds of Porzana sp. from Stranska in the Czech Republic (Monrovia) belong to “...a form larger than P. porzana, probably an extinct species.” (p. 527). No fossil taxa among the small rallids have been described from the Pleistocene deposits of Europe (Tyrberg, 1998). Three other extinct species, Zapornia palmeri (Frohawk, 1892), Z. sandwichensis (Gmelin, 1789) and Z. nigra (J. F. Miller, 1784) disappeared between 1785 and 1944 (taylor 1996). The two first species were endemic for the Hawaiian Islands, while the third one was endemic for the Tahiti Islands. Zapornia monasa (Kittlitz, 1858) from the Kosrae Island (Micronesia) disappeared by 1878. In addition, another island endemic (Z. astrictocarpus (Olson, 1973)) was described from St. Helena Island. As Olson (1977) writes “this small flightless species, along with Porzana palmeri is apparently derived from Baillon’s crake (Porzana pusilla)” (p. 356). Thus, the examined fossil specimen does not need a comparison with these species. We do not consider all insular endemic Pacific species, as well. Comparison with recent species of the genera Porzana and Zapornia Recently the genus Porzana has been split into Porzana s. s. and Zapornia, Leach 1816 (Dickinson, Porzana botunensis sp. n., a New Early Pleistocene Crake (Aves: Rallidae) from Bulgaria Table 1. Measurements (in mm) of fossil and recent rallids Species Fossil Porzana botunensis sp. n. NMNHS 301 Rallicrex polgariensis1 Recent Porzana porzana UCBL 136/1 Porzana porzana UCBL 136/2 Porzana porzana ISEA A-5157/93 Porzana porzana ISEA A-2474/71 Porzana porzana ISEA A-1848/67 Porzana porzana ISEA A-5156/93 Porzana porzana NHM 2001.17.1 Porzana porzana NHM 1957.13.1 Porzana porzana NHM 1952.2.366 Porzana porzana NHM 1976.47.1 Porzana carolina ISEA A-4667/89 Porzana carolina NHM 1986.60.14 Porzana carolina NHM 1984.77.13 Porzana carolina NHM 1984.77.12 Zapornia parva ISEA A-2538/72 Zapornia parva ISEA A-5068/92 Zapornia parva ISEA A-5155/93 Zapornia parva NHM 1999.22.1 Zapornia parva NHM 1968.4.1 Zapornia parva NHM 1952.2.365 Zapornia pusilla ISEA A-4481/87 Zapornia pusilla NHM 1996.69.12 Zapornia pusilla NHM 1978.4.1 Zapornia fusca NHM 1850.8.15.140 Rallus aquaticus ISEA A-5158/93 Rallus aquaticus ISEA A-3150/76 Rallus aquaticus ISEA A-3072/75 Rallus maculatus ISEA A-5280/94 Rallus philippensis ISEA A-4054/84 Rallus limicola ISEA A-3515/78 Rallus aquaticus UCBL 135/1 Rallus aquaticus UCBL 135/2 Rallus aquaticus NHM 1982.41.6 Crex crex UCBL 139 KR/1 Crex crex ISEA A-1611/65 Crex crex ISEA A-15150/93 Crex crex NHM 1930.3.24.59 Crex crex NHM 1930.3.24.58 Crex crex NHM 1930.3.24.56 Gallinula chloropus ISEA A-2317/70 Gallinula chloropus UCBL 140/3 Gallinula chloropus UCBL 140/4 Gallinula ventralis ISEA A-5356/94 Gallinula melanops NHM 1954.5.4 Fulica atra ISEA A-1293/63 Fulica atra UCBL 142/5 Fulica atra UCBL 142/6 Laterallus leucopyrrhus NHM 1959.10.4 Width of the diaphysis in the middle (a) Thickness of the diaphysis in the middle (b) Width of tr. m. III (c) Diameter of tr. m. III (d) 2.05 - 1.6 - 1.7 ca.2.1 2.45 - 2.2 2.4 2.2 2.1 2.2 2.2 2.5 2.0 2.2 2.1 2.0 2.2 2.3 2.4 1.9 1.7 1.8 2.0 1.9 ca. 1.9 1.8 1.9 1.8 1.9 2.4 2.3 2.6 3.1 3.0 2.9 2.4 2.6 2.7 2.8 2.7 2.8 2.7 2.8 3.8 4.8 3.4 3.7 2.9 4.0 3.2 3.9 1.9 1.8 2.2 2.0 1.8 2.0 2.1 2.0 1.7 1.8 1.8 2.0 2.2 2.1 2.2 1.6 1.4 1.5 1.6 1.6 ca. 1.9 1.5 1.5 1.4 1.9 2.3 2.2 2.8 2.4 2.7 2.4 2.2 2.4 2.1 2.6 2.3 2.3 2.1 2.4 4.0 3.2 2.8 3.1 2.6 3.6 3.5 3.7 1.6 1.8 2.1 1.9 1.7 1.9 2.0 1.8 1.9 1.8 2.2 1.8 1.9 1.9 2.6 1.4 1.5 1.4 1.5 ca. 1.7 1.5 1.4 1.5 1.8 2.2 2.0 2.3 2.4 2.6 2.8 2.4 2.2 2.2 2.4 2.4 2.4 2.3 2.3 2.2 2.7 2.8 2.5 3.3 2.3 3.4 3.5 3.6 1.6 2.6 2.6 2.6 2.6 2.8 2.9 2.7 2.4 2.7 2.5 2.9 2.8 2.8 2.4 2.0 2.5 2.3 2.3 2.1 2.1 2.1 2.5 3.2 3.1 3.6 3.2 3.6 3.1 3.4 4.2 3.0 3.2 3.3 3.3 3.1 3.1 3.4 4.4 5.4 4.0 3.7 3.5 5.0 4.9 5.6 2.3 287 Boev Z. Table 1. Continued Species Sarothrura elegans NHM 1997.34.1 Sarothrura elegans NHM 1997.34.2 Amaurornis flavirostris NHM 1992.28.3 Amaurornis flavirostris NHM 1989.19.10 Amaurornis flavirostris NHM 1966.3.3 Amaurornis flavirostris NHM 1966.3.4 Crex egregia NHM 1898.5.12.3 Canirallus kioloioloides NHM 1897.5.10.28 Porphyrio aleni NHM 1961.11.1 1 Width of the diaphysis in the middle (a) Thickness of the diaphysis in the middle (b) Width of tr. m. III (c) Diameter of tr. m. III (d) 1.7 1.7 2.8 2.6 2.2 2.4 2.3 2.8 4.6 1.6 1.4 2.1 2.2 1.8 2.0 1.9 2.4 3.3 1.4 1.4 2.1 2.1 1.9 1.9 2.4 2.7 3.0 2.0 2.2 2.9 2.8 2.8 2.6 2.8 3.6 3.1 Measurements of Jánossy (1991). He writes: “The width ... of the middle trochlea (varies) from to 2,0-2,2 mm.” (p. 22). reMSen, 2013; Del hoyo, collar 2014). Thus, two of the West Palaearctic crakes were transferred to Zapornia: Z. parva (Scopoli, 1769) and Z. pusilla (Pallas, 1776) in the Zapornini Des Murs 1860 (Old World Crakes), while P. porzana has been included in Gallinulini G.r. Gray 1840 (Coots, True Gallinules and Moorhens). As stated above, the described fossil species resembles to Porzana because of its more laterally protruded lateral edge of the distal diaphysis of tmt. above the epiphysis in the area of tr. m. IV. Size differences between P. botunensis sp. n. and the smaller Western-Palaearctic species of genus Zapornia (e. g. Z. parva, Z. pusilla) also drift it far from Zapornia. Dimensionally the specimen falls within the range of P. porzana (Table 1) but differs from that species by the more distally positioned inception of trochlea metatarsi II; shallower edges of the condyles of the trochlea metatarsi II; clearly developed edge (linea) between the facies dorsalis and facies subcutanea lateralis; deeper facies lateralis caudalis; relatively wider incissura intertrochlearis lateralis and narrower diaphysis (measurement “d”, Figs. 1, 2); from Zapornia pusilla: in the bigger size, thicker diaphysis in its distal half, absence of clear relief on the tr. m. II, less developed relief on tr. m. II and tr. m. IV, wider incissura intertrochlearis laterealis, more proximal position of foramen vasculare distale, and the almost twice wider f. m. I; from Porzana carolina: in the considerably smaller size, and the deeper fovea ligamenti collaterallis; from Zapornia fusca: in the smaller size, shallower relief of the articular surface of tr. m. II and tr. m. IV, and wider incissura intertrochlearis laterealis; from Zapornia parva: in the deeper and well-developed socket (f. l. c.) on the tr. m. II, shallower relief of the tr. m. III and tr. m. IV, wider facies dorsalis of the whole distal half of diaphysis, wider diaphysis, flater surface of facies dorsalis, more graduate but not sharp transition to the 288 Fig. 3. Distribution of the genus Porzana/Zapornia in the Pliocene – Gelasian (MN 14 – MQ 1) of Europe: 1 – Osztramos 9, Hungary (P. estramosi); 2 – S’Onix, Mallorca, Balearics, Spain (P. porzana); 3 – Beremend 16, Hungary (P. porzana); 4 – Voigtstedt, Germany (P. porzana); 5 – Stranska skala, Czech Republic (P. porzana); 6 – Beremend 17, Hungary (Z. (P.) parva); 7 – Stranska skala, Czech Republic (Porzana sp.) (1-7 after Mlíkovský, 2002); 8 – Varshets, Bulgaria (P. botunensis sp. n.) condylus medialis of tr. m. III, more asymmetrical tr. m. III, and considerably wider f. m. I. Thus, the NMNHS 301 find could not be referred to any of the recent species belonging to the genus. Discussion Garcia-r et al. (2014) state that in the Miocene and Pliocene Rallidae accomplished an increasing spatial and ecological diversification. Comparison of Nearctic and Palaearctic Regions shows that the faunistic diversity of Rallidae during the Pliocene was considerably greater in North America. FeDuccia (1968) summarises for this continent that “a mini- Porzana botunensis sp. n., a New Early Pleistocene Crake (Aves: Rallidae) from Bulgaria Fig. 4. Correlation between the measurements (in mm) “a” – width of the diaphysis in the middle, and “d” – diameter of tr. m. III in some rallids (Ref. Table 1) mum of 12 species of raillids existed during the Pliocene”, but none of genus Porzana. At the same time Bocheński (1997) lists data only on one Rallidae species, established in the Pliocene deposits from Europe. Having in mind Gallinula gigantea Tchernov, 1980, Gallinula balcanica Boev 1999, and P. botunensis sp. n., the correlation will be 12:4 for the Pliocene record (now MN 17 is placed in the Early Pleistocene). FeDuccia (1968) gives no Pliocene records of genus Porzana for North America. It seems that this genus appeared there even in the (?) Pleistocene, while in the WesternPalaearctic deposits it is found much earlier: in the Late Miocene. The Pleistocene record of Rallidae from the Palaearctic includes 14 species (Tyrberg, 1998), while in the Nearctic it consisted of 15 species (FeDuccia 1968). In the recent breeding avifaunas of both Nearctic and Palaearctic Regions the correlation between the species number of Rallids is 11:11 (taylor 1996). According to the same author only one species of the genus Porzana is spread in the Nearctic as compared to five species for the Palaearctic Region. Three species are spread in the Eastern Palaearctic and six species in South-Eastern Asia. Hence, we could determine the South-Asian region as a centre of speciation for the genus Porzana. Obviously, in Europe, the genus Pozana appeared in the Late Miocene, i.e. its chronostratigraphical distribution is MN 13 to present. The known fossil taxa (two fossil species and one fossil subspecies) were spread throughout Eastern Europe and all of them came from sites that were located in the hilly landscapes near the running water bodies. The Pliocene – early Pleistocene record of Porzana in Europe is scanty (Fig. 3). Besides the morphological differences between both species, we consider Porzana botunensis sp. n. a possible ancestor or (most probably) a fossil sister species, close to Porzana porzana. We should consider that P. botunensis sp. n. existed in MNQ 17 zone, while the oldest finds of the closest species, P. porzana, originate from the Middle Pleistocene, MNQ 22 zone (tyrBerg 1998). The summary data by this author indicates that no taxa of the genus Porzana (incl. Zapornia – Z. B.) are known from the European Quaternary before the Middle Pleistocene. The position of P. botunensis sp. n. among other species of the genus Porzana (and Zapornia) and some other small rallids (Fig. 4) shows that the Varshets specimen lays well in the scope of P. porzana, and resembles that of Z. parva. Porzana botunensis sp. n. completes the aquatic/hydrophylous avian complex of the type locality, which so far included Actitis balcanica Boev, 1998, Gallinula balcanica and Motacilla sp. (Boev, 2007). Acknowledgements: The author is very grateful to Dr. Cécile Mourer-Chauviré (UCBL), †Dr. Zygmunt Bocheński and Dr. Zbigniew Bocheński (ISEA), Dr. Robert Prys-Jones and Don Smith (Bird Group, NHM) for their valuable help during the work in their institutions. The study was sponsored by the Foundation Scientifique de Lyon et du Sud-Est (France), the Short Study Visits Programme of the Royal Society (U.K.), the National Science Fund (project No NI B-202/01.10.1992) (Bulgaria) and the NMNS. Special thanks to Dr. Nikita Zelenkov (Paleontological Institute, Russian Academy of Sciences, Moscow) and two anonymous reviewers for their critical remarks on earlier versions of the manuscript. 289 Boev Z. References Bocheński Z. 1997. List of European fossil bird species. – Acta zool. cracov., 40 (2): 293-333. Boev Z. 1995. Varhets (Western Stara Planina – Bulgaria): An example of Middle Villafranchian forest-steppe onithocoenosis. – In: Ecosystem Evolution. Internatr. Symp., Moscow, 26-30. Septr. 1995, Palaeontr. Instr., RAS, Abstracts, Moscow, 14. Boev Z. 1999. Neogene and Quaternary birds (Aves) from Bulgaria. National Museum of Natural History, Bulgarian Academy of Sciences, Sofia. D. Sci. thesis. 243 pp. + 243 pp. supplements. Boev Z. 2007. 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