Distribution, status and taxonomy of the near-threatened Black-bodied Woodpecker Dryocopus schulzi Bird Conservation International (1992) 2:253-271 Distribution, status and taxonomy of the near-threatened Black-bodied Woodpecker Dryocopus schulzi A. MADRONO NIETO and M. PEARMAN Summary Studies of records of the Black-bodied Woodpecker Dryocopus schulzi through literature searches, communications with ornithologists, personal observations and data from museum specimens show that the distribution of the species is basically limited to xero- phytic chaco woodlands in western Paraguay, northern and central Argentina, and southern Bolivia. Its western limit is defined by the transition zone between the chaco and the semi-humid montane forest on the east Andean slopes of southern Bolivia and north-west Argentina; in the extreme south it extends into the dry deciduous woodlands in the sierras of Cordoba and north-eastern San Luis. The taxonomic history of the species has been chequered; we present morphological and vocal differences from its closest relative that reaffirm its specific validity. It appears to have suffered an alarming decline over much of its former range in Argentina, and appears to survive only in two isolated population centres of significant importance, in Cordoba and adjacent San Luis and in the central part of the Paraguayan chaco. The main cause of its rarity is destruction of habitat for timber extraction and expansion of agriculture and cattle-raising. El estudio de registros del Carpintero Negro Dryocopus schulzi mediante revision biblio- grafica, comunicaciones de ornitologos, observaciones personales e information adjunta a especimenes en museos, muestra que la distribution de la especie basicamente se circunscribe al bosque chaqueno de caracter xerofilo del oeste de Paraguay, norte y centro de Argentina y sur de Bolivia. Su extremo occidental alcanza la zona de transicion entre el chaco y los bosques semihumedos montanos en las sierras orientales andinas del sur de Bolivia y del noroeste argentino; en el extremo sur se extiende hasta los bosques secos deciduos de las sierras cordobesas y del noreste de San Luis. La historia taxonomica de la especie ha sido confusa; las diferencias morfologicas y vocales que aqui se presentan y comparan con su pariente mas proximo reafirman su validez especifica. Parece haber sufrido una disminucion preocupante en la mayor parte de su distribution original en Argentina, y quedan aparentemente tan solo un par de nucleos poblacionales aislados de cierta importancia, en Cordoba y zona adyacente de San Luis y en la zona central del chaco paraguayo. La principal causa responsable de su rarefaction se debe a la destruc- tion del habitat por la explotacion de los recursos madereros y a la creciente expansion de la frontera agricola y ganadera. Introduction The Black-bodied Woodpecker Dryocopus schulzi is restricted to the central and southern chaco, in Bolivia, Paraguay and Argentina, and to the transitional subtropical forest at the low eastern edge of the Andes in southern Bolivia and north-west Argentina (see Figure 1). It was originally a candidate for inclusion A. Madrono Nieto and M. Pearman 254 Figure 1. The distribution of Dryocopus schulzi showing historical (O pre-1970), recent (• post-1970) and hybrid (•) records. Ringed areas represent the central dry Paraguayan chaco (A), humid chaco where most hybrids have been reported (B), dry chaco and transitional forest (C), and Cordoba and San Luis transitional (D); dashed line roughly separates dry chaco (to the west) and humid chaco to the east (following Hueck's 1972 map). See Appendix for key to numbered localities. in Threatened birds of the Americas (Collar et al. 1992), as its distribution and habits were poorly known (e.g. Short 1975, 1982) and most published information and correspondence (with ICBP) remarked on its scarceness (see Population below). However, after deeper investigation in the literature and further inquiries to other ornithologists with field expertise in the region, it was decided to confine it to "near-threatened" status, adopted by ICBP as a term for "birds which, while apparently not (yet) seriously in danger of global extinction, give cause for concern" (see Collar et al. 1992). However, it was also agreed that in due course the analysis already begun on the species should be completed and published, partly because it is clearly valuable to review the status of such borderline species whenever possible, and partly because this bird is a particu- larly interesting representative of the chaco fauna: any conservation initiatives in the chaco should certainly take the species into account. The Black-bodied Woodpecker 255 Methods This paper constitutes a compilation of all published and unpublished informa- tion (available to us) on the Black-bodied Woodpecker. Museums are referred to in the text by abbreviations as follows: AMNH, American Museum of Natural History; ANSP, Academy of Natural Sciences of Philadelphia; BMNH, British Museum of Natural History (Tring); IML, Instituto Miguel Lillo (San Miguel de Tucuman); MACN, Museo Argentino de Ciencias Naturales, Buenos Aires; MCZ, Museum of Comparative Zoology (Cambridge, U.S.A.); UMMZ, Univer- sity of Michigan Museum of Zoology. Part of the museum data was gathered by N. J. Collar during the early stages of preparing Threatened birds of the Amer- icas; M. LeCroy provided the information from AMNH, J. R. Navas from MACN, R. A. Paynter from MCZ, R. W. Storer from UMMZ, while we ourselves studied the specimens held in BMNH. Considerable data were collected through personal communications with ornithologists familiar with the species and from our own observations in Argentina and Paraguay. In the Appendix, records are organized within countries and provinces or departments (from north to south); coordinates are derived for every traced locality, which have been numbered and mapped in Figure 1; the coordinates are generally provided from the original source (published or not) or by refer- ence to ornithological gazetteers (Paynter et al. 1975, Paynter 1985, 1989) or DSGM (1988). Most localities in Figure 1 have been circumscribed in four general areas (A-D) in order to facilitate discussion in different parts of the text. At least one source for each record is provided, although on several occasions several sources have been included in order to facilitate further research; for this same reason if specimens exist, the relevant museum (if known) is also given (very often collecting dates and descriptions of specimens are incomplete or lacking in the literature, but information in museums has greatly compen- sated for this deficiency). Approximate altitude of the record (provided in the original source or obtained from the gazetteers) is given if available, but no attempt has been made to provide figures for localities throughout the vast low chaco plain, where elevations move gradually from below 100 m near the Para- guay river in the east to 450 m in areas at the base of the Andes in the west (Short 1975, Paynter 1985, 1989). Six study tapes of calls of Lineated Woodpecker D. lineatus from wide-ranging South American localities were compared with two available study tapes of D. schulzi from two different localities in Cordoba, Argentina (tape-recorded and published by R. Straneck; a copy is held in the British Library of Wildlife Sounds, National Sound Archive, London). Taxonomy The taxonomy of the Black-bodied Woodpecker has been the subject of much uncertainty, with various changes at both subspecific and specific level. The existence of several hybrids between lineatus and schulzi (see Appendix and below) and descriptions of invalid taxa (see below) have helped to muddle the already puzzling taxonomic position of the species. Cabanis (1883) first described the species under the genus Phloeotomus, A. Madrono Nieto and M. Pearman 256 although later generic designations of the species, or hybrids with lineatus, included: Campephilus, Ceophloeus, Dryotomus, Neophloeotomus (e.g. Dabbene 1915, Cory 1919, Peters 1948, Pergolani de Costa 1962), all of which were eventu- ally synonymized (including the lineatus and pileatus allospecies) in Dryocopus (Peters 1948; also Pergolani 1941). However, this genus (Dryocopus) was fortui- tously applied to the species by Burmeister (1861) (before it was known to science) when he referred to an immature "D. [Dryocopus] atriventris" (= Cream- backed Woodpecker Campephilus leucopogon), which, according to Cabanis (1883), was an individual belonging to the newly described species (schulzi) (see also Sclater and Hudson 1888-1889). Dabbene (1915) described a new species ("shiptoni") from Tucuman, which was identical to schulzi but for a white scapular bar; however, this taxon (although accepted by Cory 1919) was discarded by Dinelli (1931), who referred to the existence of specimens of schulzi (including paired birds) with and without this plumage feature. His opinion was supported by Mogensen (1932), who attributed this characteristic to the close relation with D. lineatus. The evidence of these two latter authors was not, however, followed by SOMA (1938) or Olrog (1959), whoxmaintained specific distinction between schulzi and shiptoni, although the latter was soon after (and finally) discarded (Pergolani de Costa 1962, Short 1975, Olrog 1979). A further taxonomic problem was introduced in 1916 by R. Dabbene, who described yet another race of schulzi, namely "major" (see Dabbene 1916, 1926). Both Mogensen (1932) and Pergolani de Costa (1962) believed this to be a good species, the former suggesting the name "Ceophloeus ater" and the latter "Dryo- copus major"; nevertheless, the distinction was not accepted by Short (1975, 1982), who considered the form in question to represent a hybridization between schulzi and lineatus. Furthermore, Peters (1926) described the subspe- cies "Dryocopus erythrops fulcitus" (type-specimen in MCZ; see also Peters 1948), which was also later considered a hybrid between lineatus and schulzi (Short 1982). A similar taxonomic problem to that of "shiptoni" (possessing white scapular bars) led to the description of "D. erythrops" (see Peters 1948), which represents lineatus without the white scapular bar, but again the specific distinction was not accepted, the form being relegated to subspecific level (i.e. D. /. erythrops) (Pergolani de Costa 1962, Short 1975, 1982, Sibley and Monroe 1990). Interest- ingly, in the south-east of the Lineated Woodpecker's range the race erythrops is distinctive (i.e. all birds lack the white bar), with a zone of overlap where both nominate lineatus and erythrops "morphs" are present (see figure 37 in Short 1975), whereas within the range of schulzi no such isolation is present inasmuch as both morphs (typical schulzi and the white-scapulared "shiptoni") can be found mixed throughout (see above) and, although southern birds are smaller than those from the northern chaco, variation is clinal (Short 1975, 1976, 1982). Despite the morphological and ecological differences between lineatus and schulzi indicated by Short (1982), that author (p.412) remarked that "whether or not it [schulzi] is specifically distinct from lineatus is a moot point". However, comparisons between the common call type of lineatus and schulzi through sona- graphic analysis tend to confirm that both represent valid taxa (see below); The Black-bodied Woodpecker 257 furthermore, the following obvious morphological differences are judged suffi- ciently important as definitely to maintain schulzi's full specific status: (a) the red crown in schulzi females extends over most of the forehead leaving only c.0.5 cm (or less) of grey feathering above the base of the bill, whereas c.1.5 cm (or more) of grey is exposed in Hneatus; (b) schulzi has pale grey or whitish ear-coverts, whereas in Hneatus this area is black or dark grey; (c) the throat in schulzi is white to dirty grey with or (usually) without fine brownish streaks (also Short 1982), whereas in Hneatus it tends to be heavily streaked giving a much darker appearance; (d) bill colour in schulzi is predominantly white, whereas in Hneatus it is typically blackish to grey; (e) nine specimens in BMNH were labelled as having "coffee" and "bright coffee" irides, whereas in Hneatus eye colour varies from white to pale yellowish orange, although the young may have brown eyes at first (Short 1982; also his plate JJ); (f) belly, flanks, vent and undertail-coverts are black in schulzi, sometimes with very fine barring, usually on the flanks and abdomen, unlike Hneatus which always shows heavy barring; (g) underwing-coverts are white in both species, but schulzi presents an irregular, usually large black patch (sometimes nearly lacking: Short 1982) on the bend of the wing, whereas in Hneatus this same area is entirely white, although occasionally with a small or, rarely, moderately sized black patch (Short 1982); (h) the rectrices in schulzi show white shafts (visible in the field and on BMNH specimens) which are dark in Hneatus. The vocalizations on both sets of {schulzi and Hneatus) recordings appear to be constant in form and duration between different localities: the loud ringing "wick wick wick" call types of schulzi and Hneatus (see Figure 2) are very similar in the quality, structure and frequency of their notes, with the fundamentals and double harmonies of Hneatus falling within the frequency ranges of those of schulzi. The vocalizations do, however, differ as follows: (a) the schulzi vocal- 1 4 - 3 - 2 - 1 - / / / ' 0.5 1.0 1.5 2.0 c CD LL 4- > 1 ' ' 1 1 1 / 1 • 1 0.5 1.0 1.5 2.0 2.5 3.0 3.5 4.0 4.5 5.0 5.5 Time (seconds) Figure 2. Sonagrams of typical calls of Dryocopus schulzi (A) and D. Hneatus (B) in the 300 Hz bandwidth. A. Madrono Nieto and M. Pearman 258 ization is delivered at four notes per second compared with the faster delivery of lineatus at six notes per second; (b) schulzi shows a broader pitch variation (see Figure 2); (c) schulzi typically delivers only five notes compared with 29 (mean from six study tapes) in lineatus. It should be noted that schulzi is capable of delivering a longer variation of this call type, although it has only been reported by one observer, and was then judged to be atypical; we would expect differences between (a) and (b) mentioned above to be constant within such a vocalization. The differences between calls in this analysis suggest that schulzi is specifically distinct from lineatus, but only a small sample for schulzi was available and the evidence presented here therefore needs further investigation. A second vocalization, "ti-chrr", with the second note being a harsh rattle, is very common in lineatus but rare in schulzi; we have been unable to make a sonagraphic comparison in the absence of a recording of this call type for schulzi. It should be noted that drumming of the two species is very similar, with lineatus giving 17.8 strokes per second (mean from five study tapes), and schulzi giving 17.6 strokes per second (mean from two study tapes), whilst duration of the drum (the same study tapes) was 1.2 seconds in schulzi compared to 1.6 seconds in lineatus, although a factor such as the state of prenuptial activity presumably precludes the possibility of a valid comparison. Whilst the morphological and vocalization differences (regardless of the ecolo- gical habitat preferences: see Short 1975, 1982; also Ecology below) are judged important enough to maintain schulzi as a valid species, the high number of hybrids recorded in areas of sympatry (area B in Figure 1), general structure of call types (see above and Figure 2) and certain phenotypic characters occasion- ally present in both species (i.e. a character not expected for schulzi but for lineatus and vice versa) serve to emphasize the common ancestral origins of these two species. Distribution The Black-bodied Woodpecker has a relatively restricted range in south-central South America, where it has been recorded from south-central Bolivia (Santa Cruz and Tarija; see Remarks 1), western Paraguay (Nueva Asuncion, Boqueron and Presidente Hayes) and north-central Argentina (Salta, Formosa, Chaco, Tucuman, Santiago del Estero, Corrientes, Santa Fe, Cordoba and San Luis) (see below; also Figure 1). Bolivia The species is only known from two localities in the central and southern parts of the country on the eastern slopes of the low Andes (see Figure 1 and Appendix). The recent record from Comarapa, Santa Cruz (record no. 1), extends the previous known range to the north by c.375 km, and it seems likely that birds will also be found in appropriate habitat in between (see Ecology). Furthermore, the record of a bird at Teniente Enciso, in the Paraguayan chaco near the Bolivian border (record no. 3), suggests that the species probably occurs in the chaco lowlands of eastern Chuquisaca and Tarija near the Paraguayan border. The Black-bodied Woodpecker 259 Paraguay All records are west of the Paraguay river where all but one (from Nueva Asuncion) are from Boqueron and Presidente Hayes departments. The record at Teniente Enciso in extreme western Paraguay suggests that the species occurs throughout the dry chaco westward to the eastern environs of the low Andes in Bolivia. The species appears to be absent from the northern chaco (Chaco department) and from south-eastern Santa Cruz department (Bolivia), where it has never been recorded (e.g. Short 1975, Remsen and Traylor 1989, Hayes et al. 1991; also in fieldwork by A.M.N.). The northern limit of the species's range in the Paraguayan chaco is yet to be ascertained. Argentina Most of the records in the country are from the western edge of the chaco in the foothills of the eastern sierras of the Andes in the provinces of Salta, Tucuman, Santiago del Estero and from the sierras of western Cordoba; however, the species also occurs to the east in the chaco lowlands reaching eastern Formosa, Chaco and Corrientes (for hybrids with Hneatus see Figure 1 and Appendix) and north-west Santa Fe (Corrientes and Santa Fe provinces are not mentioned, or included, in the distribution maps given by Narosky and Yzurieta 1987 and Canevari et al. 1991). Dabbene (1926) referred to a female collected at Santa Ana, Misiones, by F. M. Rodriguez (this specimen, according to him, had been sent to MACN), and to a second specimen taken in the "same territory" by E. Budin. We have been unable to trace the location of either of the above-mentioned specimens, which are presumably the source of other authors listing the species for that province (e.g. SOMA 1938, Peters 1948, Pereyra 1950, Olrog 1959, 1963). It is likely that there was a confusion with D. Hneatus (for which there is a large series from the same locality: see, e.g., Pergolani de Costa 1962). Later reviewers and maps of the species's distribution (see, e.g., Olrog 1979, Narosky and Yzuri- eta 1987, Canevari et al. 1991) excluded the province from its range, and there appears to have been no further mention of these two specimens from Misiones. The southernmost known locality for the species is in north-eastern San Luis province at 32°5o'S (see Appendix and Figure 1). Population Bolivia The species appears to be very rare; there are only four records from two localit- ies (in 1936 and 1991, one a hybrid: see Appendix). The record from Valle de Comarapa, Santa Cruz, much the northernmost site for the species, suggests that it may occur in the intervening region (see Figure 1), although T. A. Parker (verbally 1992) has not found it in the above-mentioned area despite extensive fieldwork there, nor did J. Fjeldsa and S. Maijer during long treks in the pre- montane zone of Chuquisaca in September-October 1991 and March 1992 (J. Fjeldsa in litt. 1992). However, as already stressed in Distribution, it is likely that the species occurs in the chaco woods of Chuquisaca near the border with A. Madrono Nieto and M. Pearman 260 Paraguay, where there is still extensive undisturbed forest (A.M.N. pers. obs.). The paucity of records from the country does not allow an assessment of past status, and it remains unclear as to whether the species has suffered a consider- able decline or was always very rare there. Paraguay Very little is known about the status of the species over a large part of the country. Most records come from the central chaco between Boqueron and Presidente Hayes departments (area A in Figure 1) within the environs of Men- nonite colonies (i.e. Filadelfia, vicinity of Lichtenau, Loma Plata, Colonia Neu- land, Orloff; see Appendix). These records suggest that the species was once and perhaps still is at least locally common (at least 13 specimens were collected in the early 1970s; also Short 1976). Neris and Colman (1991) found the species "abundant" (i.e. "found daily" in their study area; see Appendix) between April 1988 and March 1989, suggesting that it has not suffered a notable decline; however, Neris and Colman's (1991) categorization should be treated with cau- tion because daily observations in one particular locality or area may well repres- ent repetition of sightings of the same individual(s). Furthermore, A.M.N. spent a total of two months during the winter of 1989 and 1990 at site no. 10 (see Figure 1), which is very close to the locality indicated by Neris and Colman (1991), but only observed a single individual, whereas other species of Picidae were frequently encountered. It is also worth mentioning that F. E. Hayes (in lilt. 1991, 1992) spent roughly a month in the central Paraguayan chaco, where he observed two different birds on the same day twice at localities nos. 7, 8 and 9 (see Appendix). The lack of additional records from the remaining Paraguayan dry chaco (at least near the Bolivian border) presumably reflects the lack of fieldwork in this area and the species's status there remains, for the time being, undetermined. Argentina After the species was described (Cabanis 1883; see Remarks 2) Stempelmann and Schulz (1887) listed it for the province of Cordoba as a permanent resident ("not rare": Frenzel 1891), and Lillo (1902) also included it in his list of the birds of Tucuman; however neither of them gave an indication of its status. By 1910 the species had still only been recorded from the above-mentioned provinces (Dabbene 1910). Menegaux (1925) reported it for Santiago del Estero for the first time (vicinity of Icano), thus considerably extending the previous known range south-south-east into the chaco lowlands. He described nesting habits (see Ecology) and referred to pairs and groups of "five to six individuals"; further- more he indicated that the species was very rare in collections, as already noted by Dabbene (1915). Dinelli (1931) collected 11 birds at Las Termas, Santiago del Estero, between 10 and 21 September 1930; although he did not refer to the status or abundance of the species, the large number of specimens taken sug- gests that it was at least locally common in the area. Mogensen (1932) referred to the scarcity of the species in Tucuman, despite there being at least 22 specimens collected in the province between 1909 and 1928 (18 of which were taken The Black-bodied Woodpecker 261 between 1909 and 1918; see Appendix). Nores et al. (1983) judged it to be "fairly scarce" in Cordoba, an opinion shared by D. Willis (in litt. 1991); Narosky and Yzurieta (1987) considered it "rare or very difficult to find"; Canevari et al. (1991) judged it "rare" throughout its range and "poorly known", and M. Nores (in litt. 1992) believes it to be scarce although remarking that it appears always to have been so. The evidence presented above suggests that the population in the Argentine chaco (area C in Figure 1) may have declined severely, there being only four recent (post-1970) records (see Figure 1 and Appendix) compared with 30 (from 15 different sites; see Remarks 3 and Threats) historical (pre-1970) records. In the sierras of Cordoba (Grande/Comechingones) and nearby areas (e.g. north-east San Luis province, area D in Figure 1), the species has been recorded more often (although considered scarce: see above) and there are recent (post-1970) records for most known localities, in some of which (e.g. nos. 43, 51, and 46) it is regularly observed (Canevari et al. 1991, R. J. Straneck in litt. 1992; also Appendix). In this region the Black-bodied Woodpecker appears to be relatively safe in those areas where the forest remains fairly well preserved. The area B (in Figure 1), i.e. humid chaco, is considered possibly atypical habitat for the species (see below; also Short 1975), with five of the eight records there being hybrids, and the occurrence of the species in it may be only occasional: for instance, in Rio Pilcomayo National Park there were only two sightings in c. 8 years in the 1980s by a park guard (verbally to M.P.), and many ornitholo- gists visited the area without observing the species (J. C. Chebez in litt. 1992; see Remarks 4). Finally, it is worth noting the lack of records between areas A (in Paraguay) and C (mainly in north-east Salta and the western parts of For- mosa and Chaco), which may be attributable to a general lack of fieldwork in the area. Ecology The Black-bodied Woodpecker inhabits the central and southern dry chaco of Paraguay, Bolivia and Argentina, the isolated mountain range of Sierra Grande/Comechingones of western Cordoba and north-eastern San Luis, and the fringe of the east Andean sierras in Bolivia and Argentina. In the latter ecotone, the vegetation where the species occurs is a transitional gradation between the chaco and that of the high sierra, being above 1,500 m (Short 1975, 1982, Nores et al. 1983, Clarke 1991, data in this paper: Remarks 5; Figure 3). The highest elevation at which the species has been recorded is 1,800 m (in Santa Cruz, Bolivia) (Clarke 1991), but most records are below 1,000 m (see Appendix). At the above-mentioned locality (no. 1), the habitat where the spe- cies was observed corresponded to the division between semi-humid montane forest with a predominance of Alnus and Tipuana and semi-arid intermontane vegetation, where more xerophytic vegetation was dominated by cacti spp., Acacia, Prosopis, Schinopsis and Tipuana (Clarke 1991; also Remsen and Traylor 1989). The species is unrecorded north of Joaquin V. Gonzalez, Salta, in the continuing strip of montane forest (semi-humid forest or "yungas") which extends into Bolivia, where extensive fieldwork has been conducted in Calilegua National Park, Jujuy, and Baritu National Park, Salta, indicating that the species has never occurred in this ecotone. Within the humid chaco (see Figure 1), the A. Madrono Nieto and M. Pearman 2 6 2 Figure 3. Transitional woodlands at Cerro Uritorco, Sierra Grande, Cordoba, Argentina, March 1991 (photo: M. Pearman). species has rarely been recorded in Formosa (two records); at Rio Pilcomayo National Park birds were observed in an isolated woodland dominated by Pro- sopis sp. surrounded by marshland (park guard verbally to M.P.). Given that five of the seven other humid chaco records refer to hybrids (see Figure 1 and Appendix), it seems conceivable that the humid chaco is unusual habitat for the species, and further investigation on this hybridization zone needs to be conducted in order to shed more light about the status of the species there. Brief descriptions of the habitat where the species has been recorded in the dry chaco of central western Paraguay (Boqueron department) can be found in Steinbacher (1962) and Neris and Colman (1991); R. Straneck (in litt. 1992) describes the area where he has observed the species on different occasions (locality n o . 51) as dominated by Lithraea molleoides, Celtis tala and Acacia caven (see also Nores et al.'s 1983 descriptions of the vegetation of this general area). For more detailed descriptions of the vegetation throughout the species's range see, e.g., Hueck (1978), SAB (1982) and Spichiger and Ramella (1989). M. Nores (in litt. 1992) has reported that the species also appears in semi-modified areas, perhaps thus indicating a degree of adaptability to environments affected by man. There is no information about food or feeding behaviour other than that foraging occurs on trunks and major tree limbs in typical woodpecker fashion (pecking and probing, hammering, etc.) (B. M. Whitney in litt. 1991, pers. obs.). The breeding season has been given as October and November, with a moult following nesting from February to April (Short 1982); two birds collected in The Black-bodied Woodpecker 263 October in the vicinity of Lichtenau had enlarged gonads (Short 1976). The nest is drilled in the trunk of dead trees (Menegaux 1925); a pair bred in a telegraph pole by a secondary road at Alta Gratia, Cordoba (M. Nores in litt. 1992); a pair frequented a roost/nest hole in a dead tree in May 1991 (M. Sulley verbally 1992), and a bird was observed drumming on a telephone pole in March 1991 (F. R. Lambert verbally 1992). Pairs or single birds are usually reported. However, Menegaux (1925) observed groups of five or six birds (presumably a family group). Nothing is known about seasonal displacements or other questions such as territory size, clutch-size, breeding success, etc. The species appears to be a permanent resid- ent at least in Argentina and Paraguay (Nores et al. 1983, Neris and Colman 1991). Threats Bolivia The chaco habitat in Tarija and Chuquisaca is being cleared in places for water and oil prospecting (T. A. Parker verbally 1992) which, as in other parts of the chaco, is associated with human colonization and thus further deforestation. Paraguay Neris and Colman (1991) noted the increasing deforestation of the central Para- guayan chaco for agriculture and cattle-raising. This deforestation became more severe after the settlement of Mennonite farmers during the early decades of the 1900s; although they started with a subsistence economy based on agricul- ture, from the 1950s onwards they modernized their technology and thus pro- duction increased enormously, inevitably involving deforestation at an alarming rate for additional farmland, pasture and wood extraction (notably "quebracho Colorado" Schinopsis balansae). Furthermore, electricity in the Mennonite colonies (e.g. at Filadelfia, Loma Plata) is produced by wood burning in electric plants. Another reason for concern is the relatively recent (early 1980s) introduction of the "jojoba" Simmondsia chinensis, notably in the relatively pristine chaco near the Bolivian border, resulting in large areas being cleared. This plant from the Sonoran Desert of Mexico and the U.S.A. is cultivated for the production of fine oils and cosmetics. Argentina Bucher and Nores (1988) and Canevari et al. (1991) indicated that the species may be negatively affected by the destruction of its natural habitat. This is clearly reflected by the notable absence of records from the dry chaco, undoubtedly owing to the extensive logging of quebracho (Aspidosperma spp., Schinopsis balansae) and algarrobo (Prosopis spp.) for charcoal, tannins, railway sleepers and land clearance for agriculture, which has occurred on a vast scale since European colonization and continues today. Much of the land in which the species was previously recorded in area C (see Figure 1) is now deforested, accounting for the lack of recent records. A. Madrono Nieto and M. Pearman 264 Conservation Protection in the form of managed reserves is urgently required to secure the habitat where the two remaining healthiest populations occur: the dry chaco forest of west-central Paraguay (area A) - i.e. the "central Paraguayan chaco" - and the dry forest in the sierras of Cordoba (area D). Within these areas the species only receives protection in Argentina's Chancani and Copo Provincial Parks (4,920 ha and 114,250 ha respectively; protection in the latter is insufficient), both areas deserving more attention from the conservation point of view (J. C. Chebez in litt. 1992). The population in the central Paraguayan chaco is clearly under great threat (see Threats) and the creation of reserves there is of great priority before yet another population disappears. In Paraguay the species should be searched for in Teniente Enciso and Tinfunque National Parks, which are adjacent to area A (F. E. Hayes in litt. 1992). Meanwhile, ecological and population studies are needed in order to design a network of reserves that would guarantee the species's survival in the long term. Addition- ally, in the Paraguayan chaco the main Mennonite colonies desperately need electrification in order to avoid wood burning for electricity; this should be available from Itaipu Binational Dam in the Parana basin (F. E. Hayes in litt. 1992). The status of the species in Bolivia and in the Argentine chaco of Salta, western Formosa and Chaco, including its possible presence in Chaco National Park and Formosa National Reserve (J. C. Chebez in litt. 1992), needs clarifying as well as in the hybridization zone (area B in Figure 1). Remarks 1. Short (1982) erroneously referred to the species as occurring in Chuquisaca department, Bolivia; Remsen et al. (1986, 1987) published the details of this record (given to them by Short) as: "CH [Chuquisaca]: Monteagudo, 325 m, 13 May 1917 (Mus. Comp. Zool. #86762)". Paynter (1992) spotted this mistake (also present in Remsen and Traylor 1989); the specimen was collected by L. M. Dinelli in Monteagudo, Tucuman (R. A. Paynter in litt. 1992). 2. Although Cabanis (1883) only indicated that the species was observed in "central Argentine" by M. F. Schulz, the type-locality has been attributed to Tucuman (e.g. Cory 1919, Peters 1948, Pergolani de Costa 1962), while Dabbene (1915) indicated that it was discovered by Schulz in Cordoba. 3. This refers to different dates of observation without taking into account the number of birds observed on each given date. 4. Records nos. 18 and 19 are sightings and it is therefore difficult to assess whether these birds might have been hybrids as well. 5. The records in Salta, Tucuman and western Santiago del Estero (Figure 1 and Appendix) are on the western edge of the chaco and in the transitional forest with the "yungas" (see, e.g., Figure 1 in Nores and Cerana 1990, and the map in Esteban 1969: 93). The Black-bodied Woodpecker 265 Acknowledgements We would like to extend our thanks to the following persons for putting their personal observations or information/comments at our disposal: J. Fjeldsa, F. E. Hayes, P. V. Hayman, F. R. Lambert, M. Nores, T. A. Parker, R. A. Paynter, G. Pugnali, M. Sulley, B. M. Whitney and D. Yzurieta, with a special mention for J. C. Chebez and R. J. Straneck for their interest and effort in obtaining a number of records which greatly improved this paper; to R. Ranft of the British Library of Wildlife Sounds, National Sound Archive, London, for the use of audio-spectogram instrumentation and the production of sonagrams; to V. I. Mees-Balchin and R. J. Straneck for use of recorded vocalization material; to A. J. Long (ICBP) for preparing the map and adapting the sonagram; to M. LeCroy (AMNH), J. R. Navas (MACN), R. A. Paynter (MCZ) and R. W. Storer (UMMZ) for providing specimen data from their museums, and to P. R. Colston and R. P. Piys- Jones for permission to study skins in BMNH. Some of the preparation of this paper was undertaken as part of ICBP's Red Data Book programme, and it was completed with the use of ICBP's facilities and database. Appendix. Summary of the records for Black-bodied Woodpecker Dryocopus schulzi throughout its range. Country Bolivia (Santa Cruz) Bolivia (Tarija) Paraguay (Nueva Asuncion) Paraguay (Boqueron) Location/Coord./Alt. (m) (1) Cerro Picacho, Comarapa i7°53'S 64°3o'\V (1,800) (2) Villa Montes 2i°i5'S 63°3o'W (c.6oo) (3) Teniente Enciso 2i°i5'S 6i°3o'W (4) 195 km west of Puerto Casada c.22°05'S 59°45'W (5) 265 km W Puerto Casado c.22°io'S 6o°i5'W (6) Orloff, Colonia Mennonita c.22oi9'S 6o°oo'W (7) Loma Plata 22°2i'S 59°5o'W (8) South of Filadelfia 22°2l'S 6o°O2'W (9) Estancia (= Fortin) Toledo 22°2l'S 6O°2O'W (10) Estancia Campo Verde c.22°25'S 6o°3o'W Date 23.01.91 29.10.36 30.10.36 4.11.36 21.07.75 7.06.37 9.06.37 20.03.38 24.06.36 28.11.56 15.04.57 8.12.1988 8.12.1988 22.01.1988 27.07.1989 No. of birds involved * 2 *1 1 1 1 1 1 1 1 1 1 * 1 *1 * 2 • l Source Clarke 1991 Bond and Meyer de Schauensee 1942, Remsen et al. 1986, 1987 Short 1982, ANSP Remsen et al. 1986, 1987 Remsen et al. 1986, 1987 AMNH UMMZ UMMZ UMMZ Brodkorb 1937 Steinbacher 1962 Steinbacher 1962 F. E. Hayes in litt. 1991 F. E. Hayes in litt. 1991 F. E. Hayes in litt. 1991 A. Madrono Nieto A. Madrono Nieto and M. Pectrtnan 266 Appendix, (cont.) Country Paraguay (Pres. Hayes) Argentina (Salta) Argentina (Formosa) Argentina (Chaco) Location/Coord./Alt. (m) (11) 48 km west of Colonia Neuland 22°35'S 6o°3o'W (12) 50 km south of Orloff c.22°5o'S 6o°oo'W (13) vicinity of Lichtenau c.22°5o'S 59°4o'W \ (14) 70 km south of Lichtenau c.23°io'S 59°4o'W (15) "Lapachio", 110 km west of Conception (presumably Los Lapachos) c.23oi8'S 58°2o'W (16) c.6 km north-east of Joaquin V. Gonzalez 25°o6'S 64°O7'W (17) Metan 25°29'S 64°57'W (850) (18) Section Laguna Blanca, Rio Pilcomayo National Park c.25°io'S 58°2o'W (19) Guaycolec 25°59'S 58°n'W "Chaco, Argentina" (20) Castelli 25°59'S 6o°38'W (21) Napalpi 26°54'S 6o°o8'W (22) Rio de Oro (= General Vedia) 26°56'S 58°4o'W (23) rfo San Juan (possibly Riacho San Juan: Paynter i9»5)f (24) Las Palmas 27°O4'S 58°42'W Date 1988-1989 28.11.1956 27.07.1960 10.02.1970 7.06.1970 9.06.1970 10.06.1970 25.03.1971 7.10.1972 28.10.1972 4.05.1974 24.05.1974 22.07.1972 not given 04.1990 1.06.1905 1980s 1980s 21.11.1985 1988 6.06.1976 07.1924 not given 24.11.1924 19.04.1915 No. of birds involved *S 1 1 1 1 2 1 1 1 1 1 2 1 XI * l 1 * i * i • 1 1 * i 1 XI X I XI Source Neris and Colman 1991 Steinbacher 1962 Steinbacher 1962 AMNH AMNH AMNH AMNH AMNH AMNH AMNH AMNH AMNH AMNH Short 1982 Gardner and Gardner 1990 AMNH Park guard verbally Park guard verbally T. Narosky per J. C. Chebez ANSP T. Narosky per J. C. Chebez Steullet and Deautier 1946 Short 1982 Dabbene 1916, Pergolani de Costa 1962, Short 1982; MACN Peters 1926, Mogensen 1932, Short 1982; MCZ The Black-bodied Woodpecker Appendix, (cont.) 267 Country Argentina (Santiago) del Estero) Argentina (Tucuman) Location/Coord./Alt. (m) (25) Resistencia 2 / 2 7 ' S 5 8° 5 9 'W (26) Reserva Provincial Copo 26-05'S 62°oo'W (27) Las Termas 27°29'S 64°52'N (260) (28) Villa Rio Hondo 27°3 4'S 64°57'W (29) Girardet 27°37'S 62°io'W (30) Campo del Cielo 27°53'S 6i°49'W (31) Sierra de Guasay^n 28°oo'S 64°5o'W (32) Frias 28°39'S 65°O9'W (33) near Icano 28-41 'S 62°44'W "Tucuman" (34) Trancas 26-13'S 65-17'W (782) (35) Vipos 26°29'S 65°22'W (786) (36) Leales Medio (presumably c. Leales) 27°i2'S 65°i8'W (310) (37) Concepci6n 27°2o'S 65-35'W (38) Monteagudo (39) Toro Muerto 27°35'S 65°2o'W?: Paynter 1985 Date 19.07.1915 7.08.1915 11.1915 18.02.1989 11.1929 10.09.1930 15.09.1930 19.09.1930 20.09.1930 21.09.1930 8.04.1947 09.1923 23.11.1927 28.10.1980 16.11.1946 07.1903 3.08.1929 10.1926 3.04.1913 5.07.1917 15.08.1915 3.02.1909 8.07.1916 26.10.1924 28.07.1928 13.05.1917 31.10.1918 2.11.1918 4.11.1918 No. of birds involved X I X I X I *1 1 2 3 2 2 2 1 1 2 2 1 1 1 2 1 1 1 1 1 1 1 1 2 1 3 Source Peters 1926, Short 1982 Dabbene 1916, 1926, Short 1982 Dabbene 1926 D. A. G6mez per J. C. Chebez in litt. 1992 Dinelli 1931 Dinelli 1931; BMNH Dinelli 1931, Nores et al. 1991; IML Dinelli 1931, BMNH Dinelli 1931, BMNH Dinelli 1931, BMNH Nores et al. 1991 Steullet and Deautier 1946 Nores et al. 1991; MACN Nores et al. 1991 Nores et al. 1991 Menegaux 1925, Nores et al. 1991 BMNH Dinelli 1931; IML Dabbene 1915; MACN MACN AMNH AMNH MCZ MACN MACN MCZ MCZ MCZ MCZ A. Madrono Nieto and M. Pearman 268 Appendix, (cont.) Country Argentina (Corrientes) Argentina (Santa Fe) Argentina (Cordoba) LocatioiVCoord./Alt. (m) Monte Toro (untraced) (40) Laguna Pampin c.27°3o'S 58°45'W (41) Tostado 29°i4'S 6i°46'W (42) south-east of San Marcos Sierra (43) base of Cerro Uritorco Capilla del Monte 3O°48'S 64°4i'W (c. 1,000) (44) Quebrada Honda, Unquillo 3i°i4'S 64°2o'W (c.500) (45) Cordoba 3i°24'S 64°n'W (46) Parque Provincial Chancani 3i°24'S 65°27'W (620) (= Los Pocitos) (47) Quebrada de la Mermela, Chancani 3i°25'S 65°24'W (740) (48) Alta Gracia 3i°4o'S 64°26'W (49) Serranita La Rancherita 3i°4o'S 64°25'W (50) Nono 3i°46'S 65°oo'W (900) (51) Villa General Belgrano, Date 12.08.1910 08.1910? 27.02.1915 28.02.1915 03.1915? 5.11.1916 not given 26.01.1945 late 1980s 02.1990 13.03.1991 03.1991 10.05.1991 6.10.1991 17.11.1991 18.11.1991 6.06.1988 29.08.1939 currently 27.08.1980 6.01.1981 02.1989 10.07.1992 15.02.1977 not given undated 26.01- 8.02.1987 20.12.1988 No. of birds involved 1 2 1 1 2 1 1 1 n? *3 * i *i • 1 * i * i * 2 1 *2 * 2 *3 * 2 • l * 2 *1 *S *1 Source MACN Dabbene 1926 Dabbene 1926, MACN Dabbene 1926, MACN Dabbene 1926 MCZ Contreras and Contreras 1984 Giai 1950; MACN A. Johnson per J. C. Chebez in litt. 1992 G. Pugnali per J. C. Chebez in litt. 1992 M. Pearman et al. F. R. Lambert verbally 1992 M. Sulley verbally 1992 P. Hayman verbally 1992 B. M. Whitney in litt. 1991 B. M. Whitney in lift. 1991 M. Nores in litt. 1992 MCZ Canevari et al. 1991, and from information sent by R. J. Straneck in litt. 1992 D. Yzurieta per M. Nores in litt. 1982 D. Yzurieta per M. Nores in litt. 1992 J. C. Chebez et al. in litt. 1992 R. J. and M. Straneck in litt. 1992 M. Nores in litt. 1992 M. Nores in litt. 1992 D. Yzurieta verbally C. Henschke per R. J. Straneck in litt. 1992 R. J. Straneck in litt. 1992 Calamuchita 3i°59'S 64°32'W (740) 1990-1992 R. J. and M. Straneck in litt. 1992 The Black-bodied Woodpecker Appendix, (cont.) 269 Country Location/Coord./Alt. (m) No. of birds Date involved Source Argentina (San Luis) (52) Castelar, Calamuchita 18.12.1990 3i°6o'S 64°32'W (53) Parador de la Montana, 10.12.1988 Calamuchita 32°O4'S 64°38'W (1,100) (54) Embalse del Rio Tercero undated 32°i2'S 64°28'W prior 1967 (55) Chulome, Calamuchita 21.08.1980 32°i5'S 64°25'W (640) (56) near Merlo 01.1986 32°2i'S 65°O2'W (c.8oo) (56) Salto del Tabaquillo, 16.08.1992 near Merlo e.32°2i'S 65°O2'W (c.8oo) (57) La Estanzuela undated 32°5o'S 65°O3'W (840) M. Straneck per R. J. Straneck in litt. 1992 R. J. Straneck in litt. 1992 *i A. Azategui per). C. Chebez in litt. 1992 *i A. Azategui per R. J. Straneck in litt. 1992 *i G. Pugnali per J. C. Chebez in litt. 1992 *3 M. Babarskas per J. C. Chebez in litt. 1992 n? 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