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— What is Pogoniulus makawai? N.J. Collara andL. D. C. Fishpooft Qu’est-ce Pogoniulus makawaP. Le Barbion a poitrine blanche Pogoniulus makawai a ete decrit en 1965 surla base d’un seul specimen collecte dans la foret a Cryptosepalum au nord-ouest de la Zambie. Acause de l’absence d’observations ulterieures, lavalidite de l’espece aete mise en ques- tion. Les arguments suivants on ete avances: (1) il pourrait s’agir d’un individu aberrant du Barbion a croupion jaune P bilineatus, (2) bien que la localite type appartienne a un milieu tres particulier, elle n’apparait pas comme un centre d’endemisme, (3) de nombreuxobservateurs ont cherche l’espece en vain. Les auteurs presentent toutefois treize caracteristiques par lesquelles P makawaidiffere deP bilineatus un degre de difference qui, selon eux, nepeut etreattribue aune , aberration. Du reste, lestructure du milieu n’estpas dutout uniforme etles efforts pourredecou- vrirP.makawaialalocalitetypeetdanssesenvirons, bienqueconsiderables, nepeuventetrecon- siders comme exhaustifs. De vastes etendues de foret a Cryptosepalum, un habitat dans lequel il est difficile de travailler, n’ont jamais ete prospectees. Les auteurs estiment done qu’il est pre- mature de traiter P. makawai comme un synonyme. Des inventaires systematiques et complets sont necessaires, s’etendant peut-etre jusqu’aux regions limitrophes d’Angola ou en Republique Democratique du Congo. What is Pogoniulus makawaP.TheWhite-chestedTinkerbirdPogoniulus makawaiwas described in 1965 from a single specimen collected in Cryptosepalum forest in north-west Zambia. Lack of subsequent records has led to it being increasingly discounted as a valid species, because: (1) it could be an aberrant Yellow-rumpedTinkerbird P bilineatus, (2) although Cryptosepalum forest is averydistinctive habitat, the type locality does not appear to be in a centre ofendemism, and (3) many people have since searched for it without success. However, we find 13 separate char- acters by which it diverges from P. bilineatus, a degree of difference which we feel cannot be ascribed to aberration. Moreover, the habitat ofthe area is byno means uniform; and the efforts to rediscover P. makawai in and around its type locality, while considerable, cannot be regarded as exhaustive, particularly since large areas ofCryptosepalum forest, extremely difficult habitat in which to work, have never been visited. Assigning P. makawai to synonymy is, we feel, prema- DR ture; systematic and comprehensive surveys, perhaps into adjacent Angola or Congo, are needed. From the very moment of its naming, the ing conviction by two pairs of authorities, & White-chestedTinkerbirdPogoniulusmakawai Dowsett Dowsett-Lemaire (1980, 1993) and & has been haunted bydoubt overits taxonomicsta- Short Horne (1985, 1988, 2001, 2002). As a tus. The paper in whi—ch Benson & Irwin (1965a) result, the species was not reco&gnised by Sibley & described the species taken by their remarkable Monroe (1990), Dowsett Forbes-Watson & collector Jali Makawa, in whose honour it was (1993), Aspinwall Beel (1998) or Dickinson named, in an areaofCryptosepa—lum forestnorth of (2003). In the face of this substantial scepticism ‘Mayau’ in north-west Zambia was immediately on the part oftwo highlyauthoritativeworld lists, followed by a comment (Goodwin 1965) which one equally authoritative African list, and the postulated the notion that it might, in fact, be an Handbook of the Birds of the World, BirdLife aberrant Yellow-rumped Tinkerbird 7? bilineatus. International, having treated makawaias a threat- & & Owing to the subsequent inability of anyone to ened species (Collar Stuart 1985, Collar — confirm the existence ofP makawai a comment Andrew 1988, Collar etal. 1994), has since 2000 about ‘fresh material’ in Fjeldsa (2003) proving to opted to regard it—as Data Deficient (BirdLife have been unfounded (J. Fjeldsa in litt. 2005) International 200—0) meaning that its taxonomic this possibility has been entertained with increas- status is unclear although the species’ was still & 18-BullABCVol13No 1(2006) WhatisPogoniulusmakawaP: Collar Fishpool — used to help define an Important Bird Area in to pale yellow on the lower chest (throat to belly Zambia (Leonard 2001, 2005). pale whitish grey in bilineatus belly with a slight & , Short Horne (1988) retained it as a species greenish tinge); (6) lower breast to belly lacking & (with conside—rable reluctance), as did Sinclair greenish tinge; (7) central belly black (no such Ryan (2003) whose report that the voice of mark in bilineatus); (8) entire underparts below makawai is ‘subtly different from Yellow-rumped breastwith pale blackish ‘shadow-barring’ (absent Tinkerbird’ is presumably based on the comment in bilineatus); (9) underside ofthe bend ofwing & inAspinwall Beel (—1998) that the voices ofthe black, not white; (10) tibial feathering more suf- two taxa ‘may differ’ but the trend in general fused black; (11) bases offeathers on mantle and suggests that because ofthis taxonomic uncertain- underparts pale (darkin bilineatus); (12) bill heav- tyP. makawaiwill steadily disappear as a target of ier, more arched and less conical, with cutting ornithological interest and investigation, and edges of the upper mandible flared around the hence ofconservation activity. This is in spite of gape; (13) black bill whitish basally and from the two rather strong declarations in favour of nostrils to halfway along the cutting edges (all makawai as a good species, one by the late C. W. black in bilineatus); (14) rictal bristles at the level Benson in a personal communication to NJC in ofgreatest development found in any individuals & Collar Stuart (1985: 355), inwhich its ‘validity of bilineatus (15) legs and feet markedly paler; ; asaspecieshasbeenmostemphaticallyreasserted’, (16) toes and claws ‘equally pallid’ as the legs and and the other by G. R. Graves, also to NJC, feet; and (17) legs slightlylongerandmorerobust. reported in Collar & Rudyanto (2003: 107-108), From our own examination we would make in which ‘following a preliminary (two-hour) the following comments and qualifications on inspection of the type, the White-chested these numbered characters: the wing fringes (3) Tinkerbird seems likely to prove a good species’. are barely perceptibly paler but unquestionably Neither of these judgements was published in a narrower than in bilineatus in contrast to the ; place where much notice would be take—n ofit glossy, inky black ofthe rest ofthe plumage, the norperhapswasthepleabyIrwin (2003) and40 chin (4) is a matt greyish black; the central belly years after the species was named we feel the time patch (7) is aratherirregularsmudge; theshadow- has come to examine the evidence afresh, and to barring (8) is actually throughout the underparts, weigh more carefully the case for and against even on the creamy-white throat, but so slight as makawaias a taxonomic entity. to be virtually invisible on the specimen when held at arm’s length; the gape-flange swelling (12) The case for maynot be greater than in some bilineatus and as , The case for has hardly been made since the first Goodwin (1965) pointed out, some allopatric description. The only subsequent arguments in bilineatus have bills that match makawai in size; favour are the two personal judgements just quot- the rictal bristles (14) are likewise barely different ed, neitherofwhich comeswith anydetail to sup- from those on bilineatus the continuous col- ; port the conviction. The first thing to be done, oration of the legs, feet, toes and claws (15, 16) therefore, is simply to list out, as clearly as possi- form a single character; the legs are not longer ble, the diagnostic features ofP makawai as they than bilineatus and if they are more robust (17) & emerge in Benson Irwin’s (1965a) comparison this is too slight and unquantifiable a feature to with P bilineatus (not all ofwhich are explicitly allow. Thus we would say that the diagnosis of indicated as distinguishing marks, but which our makawai rests on characters 1-11, 13 and 15 comparison of text and specimens suggests was above, making 13 features in all. their intention): (1) white supraorbital stripelack- Thereis onepossiblefurtherfeaturein Benson ing; (2) whitelinebelowtheear-coverts onlycom- & Irwin (1965a): the white streak below the ear- mencing behind the gape, not running below the coverts is ambiguously described as ‘joining with eye in a continuous band from over the bill; (3) the pale under parts’ but then ‘from which it is yellowfringes to the secondaries andwing-coverts separated bya blackmalarstreak’. On balance, we paler, possibly narrower; (4) chin black, flecked interpretthis to mean thatthestreakiscontinuous centrally with white (chin whitish in bilineatus); with thewhite neck, but the illustration accompa- (5) throat and upper breast creamy white, fading nying the description clearly shows the opposite, WhatisPogoniulusmakawai?: Collar&Fishpool BullABCVol13No 1(2006)-19 with both bilineatus and makawai having this strikinglyin colour pattern than do bilineatusand lower facial stripe entirely enclosed by black. In Yellow-throated Tinkerbird P. subsulphureus and , partial contrast, the illustrations ofthe two taxa in pointed to the sharp difference in facial pattern of Short & Horne (2001) and in Sinclair & Ryan makawai and bilineatus when viewed front-on (a & (2003) show this streak meeting the pale under- feature illustrated by Benson Irwin), remarking parts in bilineatus but being enclosed by black in that ‘this difference could function as an isolating makawai as if this is a distinct character differ- mechanism as there is abundant circumstantial , & ence; and the illustrations in Short Horne evidencethatthecolorationoftheheadandupper (2002), which omit makawai, again depict bilin- breast ofbirds is often ofprimary significance in eatuswith a streakcontinuous with the pale collar this respect’. Goodwin thus concluded ‘that (although the position ofthe painted birds makes makawai is best considered as a new species, at thisveryeasytomiss). However, carefulinspection least provisionally’, and both Mayr (1971) and of the type of makawai reveals that there is no Snow (1978) followed thisjudgement, the former essential difference between it and bilineatus in adding a plea for comparative studies ofthe calls, this regard, bothtaxahavinganarrowlineofblack the latter mistakenly referring to Yellow-throated that in some positions appears to isolate thewhite Tinkerbird P. subsulphureus instead of Yellow- cheek-stripe and in others is broken by it. fronted Tinkerbird P chrysoconus as the third & Photographs in Short Horne (2002: 139) and Pogoniulusin the area. & in Ginn etal. (1989: 398) show both conditions Dowsett Dowsett-Lemaire (1980), in their in bilineatus. first of two brief comments on makawai took , much the same line as Goodwin, but, writing 15 The case against years later, pointed out that ‘several visits to the Ofwhat, then, does the case against consist? As type locality have failed to produce any further noted, there are three independent sources of evidence, and in particular no unusual Pogoniulus doubt: (a) Goodwin (1965), (b) Dowsett & vocalisations have been heard.’ After a further 13 & Dowsett-Lemaire (1980, 1993), and (c) Short years their patience had worn thinner: ‘investiga- Horne (1985, 1988, 2001, 2002). In reality, how- tions by a number ofobservers in north-western ever, Goodwin (1965) only very tentatively sug- Zambiahavefailedto rediscoverit... As anticipat- gested that ‘the possibility ofits being an aberrant ed by Dowsett & Dowsett-Lemaire (1980), we individual of P bilineatus cannot be entirely now believe it is no longer justified to recognise excluded’, and most of his commentary was makawai as other than an aberrant P. bilineatus weighted against this notion. He pointed out that (Goodwin 1965)’ (Dowsett & Dowsett-Lemaire makawai shows greater melanism on the head, 1993). underwing and central belly than bilineatus and Short & Horne were always unconvinced. , less melanism on the remaining belly area and ‘Despite intensive searches in western Zambia, breast, admitting that ‘it would be most unusual, they wrote, makawai ‘remains known from but but not unprecedented, for an aberrant individual one specimen’; and because that specimen ‘comes to have more melanin than normal in some areas from no distinctive habitat or area ofendemism, and less elsewhere’. He also pointed out that the and rather closely resembles P. bilineatus we are , greater curvature ofthe culmen and width ofthe inclined to regard itas averyaberrantspecimen of bill ofmakawaiare ‘not in themselves ofgreat sig- bilineatus (Short & Horne 1985). Three years nificance’, given thatsomeformsofbilineatushave later, in TheBirdsofAfrica (Short & Horne 1988) bills that approach and even match it in these they allowed the species an entry but were pro- characters; butheacknowledged thatspecimens of foundlysceptical: bilineatus (race mfumbiri) from near the type Status highly uncertain. Only 1 bird found, locality ofmakawai all have more slender, conical despite repeated searches... No ‘odd’ tinker- bills, suggesting some ecological separation, and bird calls have been heard or unusual indi- again admitted that ‘it would certainly be surpris- viduals seen... at type locality... P makawai ing ifan aberrantly coloured individual happened couldprovetobeanaberrantYellow-rumped also to have a slightly aberrant bill’. He then Tinkerbird, ifits distinctive features are sim- observed that makawai and bilineatus differ more ple melanism. 20-BullABCVol13No 1(2006) WhatisPogoniulusmakawai?: Collar&Fishpool — —— By the start ofthis century, in their book on simply too much to be ascribed to aberration. In barbets, their view had hardened further: they particular, the redistribution of colour pattern could nowpoint to the failure of‘three decades of the blackchin, the missingwhite supraorbital and searching by various ornithologists and bird- supraloral bands (but the retained white cheek- watchers’, and indeed theydid so twice, mention- stripe), theblackbellypatch, thepart-palebilland ingagain the ‘numerous searches’ in the ‘relatively all-pale legs, the whitish dorsal underfeathering non-distinctive habitat in which it was found’ is entirely uncharacteristic ofaberrant individuals & (Short Horne 2001). Thus makawai is ‘almost (although it is ofcourse somewhat problematic to certainly a very aberrant individual’ of bilineatus, speak ofwhat is typical of atypicality); certainly although in the caption to their illustration ofit nothing in the entry ‘Plumage, abnormal’ in & they described it as a ‘morph’ (which is averydif- Campbell Lack (1983) indicates otherwise, and ferent biological category). Even so, they gave it a we can think ofno comparable case where so dis- separate account ‘because there seems to be no tinctive a specimen has been disallowed taxonom- simple genetic explanation for all of its distinct icvalidity. features, e.g. melanism would account for some Moreover, since Goodwin’s time of writing features, butnotthelackofyellowandgreybelow, very considerable advances have been made in & nor the heavy bill found in this male’ (Short understanding the genetic basis ofblack plumage Horne2001). On theotherhand, onlyayearlater inbirds (reviewedbyMundy2005). In thelightof they remarked that makawai ‘is now generally these, the probability ofa melanin-related muta- accepted as representing an odd variant of the tionaccountingforthisdivergencedeserves recon- & Yellow-rumped Tinkerbird’ (Short Horne sideration. Across awide range oftaxa intraspecif- 2002: 143),withasimilarcommentunderthelat- ic polymorphisms in melanin-based colours have & ter species (Short Horne 2002: 184). repeatedlybeen found to be associatedwith varia- The case against makawai therefore depends tion in a single gene (MC1R), but these typically on the following points: (1) that it could be an involve a consistent increase in the extent of aberrant bilineatus', (2) thatthe type specimenwas melanised feathers, rather than the simultaneous obtainedin an areabelieved to be undifferentiated darkening and lightening seen in P makawai (as by habitat or by endemism; and (3) that searches anticipated by Goodwin). Moreover, such muta- have failed to find it or even to detect any tions are typically not associated with simultane- unknown Pogoniulus calls, with emph—asis variably ous side-effects on othertraits, such as other com- placed on the—number of searches ‘intensive’, ponents of morphology. Both of these points ‘repeated’, etc.—and simply the length oftime would tend to imply that the morphological dif- ‘three decades’ without renewed contact (Snow ferences between makawaiand bilineatusarehigh- [1978] stated, forexample: ‘All attempts to obtain ly unlikely to have arisen as a consequence of a further specimens have so far proved unavailing’). one-off mutation generating a single aberrant These three objections need to be examined in individual. turn. 2. Is the type locality undifferentiated by habitat or — The case against examined endemism? That the type of makawai comes 1. Could —it be an aberrant Yellow-rumped from an area oflow endemism and from a wide- Tinkerbird? The possibilityofmakawaibeingan spread habitat is not in serious dispute. aberrant bilineatus seems to us to have been fairly Nevertheless, Benson & Irwin (1965a) pointed well undermined by Goodwin (1963) even as he out that Cryptosepalum forest in this area could be raised it. He admitted that aberrant specimens seen as ‘an evolutionary centre’ given the presence thatareboth moreandless melanistic than typical there of‘such distinctive forms’ as the red-necked birds are highly unusual; and he further admitted race ofCrested Guineafowl Guttera edouardikath- that for any such specimen also to be aberrant in leenae (although this is synonymised in Crowe et bill morphology would compound the degree of al. 1986), plus Margaret’s Batis Batis margaritae anomaly. Apart from this, we reg—ard the evidence kathleenaeand, ‘inthispartofits range’, Gorgeous in Benson & Irwin’s description amountin—g in Bush-shrike Telephorus viridis. Benson & Irwin our judgement to 13 points of divergence as (1965b) and Benson et al. (1971) expanded on WhatisPogoniulusmakawai?: Collar&Fishpool BullABCVol13No 1(2006)-21 ? this, indicatingthatthefirstandthirdofthesetaxa that very considerable endeavours have gone into are known in Zambia only from this area of the quest for makawai. But what is the truth of & Cryptosepalum, which Irwin (2003) stressed ‘can this? Benson Irwin (1965b) were the first to hardly be described as “non-distinctive”’. T. B. report on a newsearch. The specimenwas collect- Oatley(in lift. 2003) agrees: ‘One needs to lookat ed on 6 September 1964 during a four-day the region, notjust the Cryptosepalum forest, and prospection ofthe area (3-7 September), and the Macronyx grimwoodi [Grimwood’s Longclaw] area was revisited for five days, 8-12 November (and, ifI remember rightly, some butterflies) can 1964, when ‘every effort was made to find the then be added to the list of local endemics.’ species again, but completely without success’. Benson & Irwin (1963b) offered the following The following year, in May 1965, Oatley (1969) scenario: spent three weeks in north-west Zambia, explicit- Pogoniulus makawai is according to present lyin order to document theavifaunabetterand to knowledge endemic to Cryptosepalum... The discover more about makawai', he camped near ancestral population may have been wide- ‘Mayau’ on 2-8 May, at a time when tinkerbirds spread and plentiful in a former more exten- were breeding, butheard no unusual calls, observ- sive area of Cryptosepalum. Thereafter, as ingthat ‘possiblythesingleknown specimenwas a recently as 12,000 years ago, according to vagrant from some other locality, perhaps farther Moreau, adrierperiodensued, duringwhich west in eastern Angola (and speculating that the this population may have become isolated, heavier bill might be less important in feeding and speciated into makawai. Subsequently, ecology than in hewing nesting cavities in harder under a moister, modern regime, bilineatus timber than that encountered by bilineatus). has perhaps reinvaded the Cryptosepalum. It Fifteen years after Oatley’s endeavour, Dowsett & may be in active and successful competition Dowsett-Lemaire (1980) reportedthat ‘severalvis- with makawai, which may before long its’ to thetypelocalityhaddrawn blank.Themost become extinct. important of these was by R. Dowsett with J. J. Whether or not it is plausible that makawai Makawa himselfin 1973, when they spent 9-20 speciated as recently as 12,000 years ago, this August at Mayau and collected ten bilineatus explanation forits raritymakes considerablesense. (Dowsett 1973, R. J. Dowsett in litt. 2005). The It is not necessarilythe case, however, that there is only other publication on Cryptosepalum birds in direct competition between makawai on the one north-westZambia, byBowen (1980),wasastudy side and bilineatus and, indeed, the syntopic which did not involve ‘Mayau’, although one of Yellow-fronted Tinkerbird P. chrysoconus on the the two sites surveyed was deemed very like other; rather, one might expect makawai with its ‘Mayau’, since it produced a bird list very similar more voluminous bill to occupy a feeding niche to that in Benson & Irwin (1965b). Bowen, like thatallows its co-existencewith these closelyrelat- Oatley, was constantly alert for novel tinkerbird ed species. Benson et al. (1971) made this point, calls but heard none, and speculated ‘that and F. Dowsett-Lemaire {in lift. 2005)—has point- makawaidoes not extend as far north as the areas ed out that some sp—ecies ofPogoniulus including I covered’. Altogether, therefore, the evidence indeed chrysoconus are specialists on mistletoe appears to be that relatively little work has been berries (Loranthaceae and Viscaceae) (see done in the area ofthe kind thatwould be appro- Dowsett-Lemaire 1988), so some kind ofspecial- priate to a serious endeavour to rediscover the ism in makawai would seem likely to explain the species. co-occurrence ofthree congeners. Thus A. Scott, director of the Wildlife J. — Conservation Society of Zambia, wrote to J. FT 3. Has it been exhaustively searchedfor Finally, Fanshawe at BirdLife International in 1989 to there is the number oftimes that the type locality report that ‘no-one has made any attempt to find and nearbyareas ofCryptosepalum have been visit- [P. makawai\ since Bob Dowsett... in the early ed with no evidence of makawai being found. 1970s’. From 1989 things evidently changed From some ofthe language used by those consid- somewhat. R. J. Dowsett (in litt. 2005) himself ering the issue (reference to ‘intensive’ surveys and has very helpfully enumerated the observers who ‘three decades’ ofsearching), it is easy to assume havebeen to thetypelocality, andhesuggeststhat, 22-BullABCVol13No 1(2006) WhatisPogoniulusmakawai?: Collar&Fishpool in all, more than two man-months have been always been Golden-rumped and Yellow- spent there bycompetent fieldworkers: fronted. No-one has been trying to call up each and every tinkerbird or concentrate Dylan Aspinwall (several times); Carl Beel fully on them. It is definitely true that no (incl. 1-3 Sep 1993); Clide Carter (several hardeffortwasmadetoproveordisprovethe visits), with Nigel Hunter (5-6 Oct. 1989); existence of P. makawai. Nearly all efforts Pete Leonard (i&ncl. Aug. 1996); Jorg were near the type locality. It might just be Mellenthin (Feb. Aug. 2000, also 2002); possible that the bird is not in prime habitat Bob Payne; Bob Stjernstedt (incl. 14 Apr. (anymore?) at that spot, especially as more 1974); Paul van Daele (incl. Sep. 1998). people have moved into the areaandcutlots Details of other visits are unknown, but of trees and burnt undergrowth. There are everyone who has lived in Zambia for any large tracts of Cryptosepalum remaining, but length oftime hasvisited the area (and some difficult ofaccess and unvisited. visitingbirders), hopingtobecomefamous. I have also passed by Mayau briefly on other Slightly more recently, P. M. Leonard (in litt. occasions. In addition, Fran^oise [Dowsett- 2004) has commented as follows: Lemaire] and I spent 12 days camped a bit Very few serious expeditions have taken further south in the neighbouring West place. Perhaps Terry Oatley’s in the 60s was L&unDgoawsNeattti-oLneamlaPiarrek1i9n7N8o)v,.wi1t9h78no(Dsoiwgsnetotf hthaevemossttayseedriomuos,rebutthIanknaowcooufpvleeryoffewniwghhtos anyunusual tinkerbird. before being driven away by sweat-bees and C. Beel (in litt. 2002) informed us that ‘Clide truly impenetrable jungle. I don’t think the Carter, DylanAspinwallandBobStjernstedttried’ searching has been thorough at all. to find it at some stage, although the sadly late Everybody tags it onto a Mwinilunga trip as Aspinwall (in litt. 1981 and 1994) did not men- a token gesture, but it needs a month of hardcore canopyscanning. tion such attempts and indeed in 1994 reported that he had no immediate plans to investigate the This testimony is important: it suggests that a species; by that stage he was inclining to the view relatively rare bird of the canopy could easily go that makawai was most probably an aberrant undetected in such dense and unwelcoming habi- bilineatus, which is, as&already noted, how it was tat (Beel stressed the need to provide plenty of treated in Aspinwall Beel (1998). Meanwhile wateronavisit, andtoprepareforthesweat-bees). Beel himselfbelieved that only he, P. M. Leonard Ifin fact there is some subtle habit—at selection by and P. van Daele had visited the area in recent makawaiwithin the Cryptosepalum for ex—ample, & years for any time, although Ryan Cassidy perhaps in tallergrowth alongwatercourses then (2003) paid a briefvisit, and bird tours stop there the lack of records since 1964 might be all the to find Gorgeous Bush-shrike. Beel (in litt.) more understandable. continued: T. B. Oatley (in litt. 2005) is surprised at Beel and Leonard’s comments about the impenetrabili- Nearly all of us have spent time along the ty of the habitat, and offers the following main Mwinilunga-Kabompo road. It is not easy to enter the forest away from this road. reflexions: There is (was?) a narrow track to a game In 1965, the mavunda was not Very dense camp on the edge ofWest Lunga National and tangled’ or ‘hard to getaround in’ and it Parkwhich starts at Mayau. I tried to follow certainlywas not ‘trulyimpenetrablejungle’. it, lost my car antenna and nearly my mir- Infact, onceonegotthroughthedenseedge- rors, then turned back. This track crosses a effect road fringe, one could walk around vast block of Cryptosepalum where no-one over a fairly open forest floor with scattered has tried to look. Cryptosepalum itselfis very thickets of undergrowth [R. J. Dowsett in hard to get around in; it is very dense and litt. 2005 reports similar conditions in tangled. This makes it very difficult to get a 1973]. It sounds to me as though there has good look at a tinkerbird even a short dis- been heavy exploitation ofthe taller timber tanceintotheforest. I thinknoneofusspent (resulting in dense secondary growth) in the morethanaday, maybetwo in thearea.Very area,whichliesdistantfromdistrictadminis- fewtinkerbirds can be seen. Those seen have trative centres and is probably seldom WhatisPogoniulusmakawai?: Collar&Fishpool BullABCVol13No 1(2006)-23 policed by forestry officers. I note that Beel With respect to this last point, it is worth noting explicitly refers to lots of tree cutting... that the type locality is also right on the edge of Thinking over the possibilities, we concen- the range of bilineatus in Central Africa (see the trated most of our searching in the maps in Snow 1978, Short & Horne 1988), so it Cryptosepalum forest patches, assuming that would seem plausible that makawai may prove to makawai was a forest canopy bird. But it be a replacement ofbilineatus in slightly different need not have been. The area where it was habitats in this area. collectedwas amosaic ofmiombowoodland Certainly a concerted, systematic endeavour and mavunda forest, and bird parties from over several months is likely to be required either the woodland regularly entered the canopies to relocate makawaiwithin the general area ofthe ofthe forest patches that were in their path. Another point that arises is that September, ltoycpael alovciaflaiutnyao—rbtuotdeivsecnoutnhtenitwperocpoeurlldynfortocmoutnh-e when Jali collected the bird, is the early springmonthwhen birds (especiallyfloaters) tenance abandoning the species to synonymy move around a lot looking for mates or without much more work further west or north. vacancies in the breeding population. (A hybrid origin of makawai seems, incidentally, Sept/Oct is also the time when many altitu- never to have been mooted, but ifa hybrid shows dinal and otherAfrotropical migrants are on adegreeofch—aracterintermediacyand bilineatusis the move, the time when one can find odd one parent—both of which are reasonable birds brieflysojourningin atypical habitats! assumptions noplausiblecandidatefortheother parentpresentsitself; onthispointtheexplanation To this M. P. S. Irwin [in litt. 2005) adds some at once founders. Another consideration is that further thoughts: the type specimen of makawai might yet possess DNA With long experience of collecting in the sufficient to test against bilineatus, w—hich field, it is extremely difficult to say that possiblywould resolve the debate atastroke but something is not there or has been missed. wecandonomorethanencouragetheexploration Move camp a mile or two into aslightlydif- ofthis notion.) ferent habitat and there will be a complete Meanwhile, from our study ofthe type speci- change in the species one is likely to collect. men in the Natural History Museum, Tring, UK, Allcollecting [atMayau] wasdonebackfrom we can confirm 13 points of distinction from P. the river ofthat name where the forest was bilineatus mfumbiri indicated by Benson & Irwin densest. But what about somewhere miles (1965a) as enumerated above (some ofthem evi- away? And look at the Cryptosepalum in dent on the accompanying plates). The specimen Angola, which seems particularly dark and (BMNH 1964.33.1) was an adult breeding male, dense and quite unlike that at Mayau. testes 9.0 x 6.5 and 7x6 mm, wing 56 mm, tail 32 mm, tarsus 15 mm, culmen from base ofskull Conclusion mm & 13 (Benson Irwin 1965a); its skull was All ofthese factors lead us to the conclusion that fullyossified (BMNH label data). Itwas collected there is no firm basis yet to discount Pogoniulus ‘high up’ in the canopyofCryptosepalum forest on makawai as a good species. From this it must fol- Kalahari sand on 6 September 1964, four miles low that either (a) it is very uncommon through- north of Mayau, Kabompo District, Zambia, at out Cryptosepalum forest in north-westZambia, or roughly 12°42’S 24°16’E (Benson & Irwin (b) it is restricted to a relatively uncommon habi- 1965a). The geographic component ofthis infor- tatwithin oradjacentto Cryptosepalum (ifthetype mation was adjusted by Dowsett (1980): was taken on the Mayau River, then perhaps itwas in or near riverine forest, not Cryptosepalum), or ...because ofthe importance ofdiscovering t(ch)is, aasreOaatflreoym(s1o9m69e)whsaptecudlifafteerde,ntithiasbaitsattrsagfgulretrheirn afbulrethteor psupbelciismhenass aocfcutrhiastespaectiyepse-iltociaslidteysira-s west in near-adjacentAngola, averylittle explored possible. There is no locality named Mayau, area, to which we would add the southern butthereisariverofthatname (orMayowo) and a plain. The specimen was collected Democratic Republic ofCongo, close to the bor- about 6 km north ofwhere the track from derofwhich the type localityofmakawaialso lies. 24-BullABCVol13No 1(2006) WhatisPogoniulusmakawai?: Collar&Fishpool Figures 1-3.ThetypeandonlyspecimenofPogoniulus makawai(BMNH 1964.33.1, above), anadultmale, takenneartheMayauRiverin north-westZambia, September 1964,withanadultmaleP. bilineatus (BMNH 1964.33.2, below) takenatthesamesitethree daysearlier. Characters ofmakawaivisiblehereare: no whitesupraorbitalstripe;whitelinebelowear-coverts commencingonlybehindgape;yellowedgestosecond- ariesandwing-covertsnarrower; chinblack, fleckedcen- trallywhite; throatandupperchestcreamy-white, fading topaleyellowonlowerchest; lowerchesttobellywith- outgreenishtinge; central bellyblack;whole underparts belowchestwithblackish ‘shadow-barring’; tibia!feath- eringmoresuffusedblack; blackbillwhitishbasallyand fromnostrils tohalfwayalongcuttingedges; legsand feetmarkedlypale. Othercharactersincludeundersideof bendofwingblack, notwhite; basesoffeathers onman- tleandunderpartslight, notdark. L’uniquespecimen (lespecimentype) dePogoniulus makawai(BMNH 1964.33.1, enhaut), unmaleadulte, collectepres delaMayau, Zambiedunord-ouest, sep- tembre 1964, avecunmaleadulteP. bilineatus(BMNH 1964.33.2, enbas) collectealamemelocalitetroisjours auparavant. Lescaracteristiquessuivantesde makawai peuventetrenotees: absencedetraitsupraorbitalblanc; traitblancsouslajouecommen^antseulementenarriere delacommissure; remigessecondairesetcouvertures alairesaliseresjaunesplusetroits; menton noir, tachete deblancaucentre; gorgeethautdelapoitrineblanc- creme, devenantjaunepalesurlebas delapoitrine; absencedeteinteverdatresurlebasdelapoitrineetle ventre; milieuduventre noir;ventrebarredesombre; plumes dutibiateinteesdavantagedenoir; beenoir, avec labasedes mandibulesblanchatredepuis lescommissures jusqu’alamoitiedubee; pattesnettementpales. D’autres caracterescomprennent: dessous delacourbedel’aile noir, nonpasblanc; basedesplumes du manteauetdes partiesinferieuresclaire, passombre. CreditforFigure 1: N.J. Collar (©NaturalHistory Museum) CreditforFigures 2-3: C. N. Spottiswoode (©Natural ' 1 HistoryMuseum) 3 Mwinilungato Kabompo crosses theMayau; or not, spreading out methodically into the sur- m this is at an altitude of 1,150 at about rounding areas, with a particular interest in sam- 12°42’S 24°15’E. pling any slight modifications of Cryptosepalum As precisely as possible, it is at this site that a forest caused by water or other features, taking systematic long-term search for the species should special note ofmistletoe taxa and abundance, and commence,whetherthehabitathasbeenmodified with an eyeon the fact that atthis locality bilinea- WhatisPogoniulusmakawai?: Collar&Fishpool BullABCVol13No 1(2006)-25 — tusis, apparently, at the southern edge ofits range Benson, C. W. 6c Irwin, M. P. S. 1965a.Anewspecies in the country. oftinker-barbet from Northern Rhodesia. Bull. Br. Therewas a timewhen the failure ofornithol- Omithol. Cl 85: 5-9. ogists to locate the Red-tailed NewtoniaNewtonia Benson, C. W. 6c Irwin, M. P. S. 1965b. The birds of fanovanae was attributed to the type and only Cryptosepalum forests, Zambia. Arnoldia Rhod. specimen being an aberrant Red-tailed Vanga 1(28): 1-12. Calicalicus madagascariensis (see Collar & Stuart Benson, C. W., Brooke, R. K., Dowsett, R.J. & Irwin, 1985). This view was, incidentally, strongly M. P. S. 1971. TheBirds ofZambia. London, UK: opposed by none other than C. W. Benson Collins. (Benson etal. 1977), but it took years before the Benson, C. W., Colebrook-Robjent, J. F. R. 6c Williams, A. 1977. Contribution a fornithologie increased ornithological interest in Madagascar & de Madagascar. Oiseau R.F.O. 47: 41-64. from the early 1980s yielded aconclusive result BirdLife International. 2000. Threatened Birds ofthe i(nGdoeoedd,mantwo& vSicrhtuuallelnybersgimu1l9t9a1n;eoaulsso rEesvualntss BoweWno,rldP..SCt.aJm.b1r9i8d0g.e,ThUeK:birBdisrdoLfifCeryIpnttoesrneaptailounmalf.orest 1991). The rediscoveries of the Yellow-throated nearMwinilunga. Bull Zambian Omithol. Soc. 12: Serin Serinus flavigula and Sao Tome Grosbeak 48-54. Neospiza concolor, each after over 100 years of Campbell, B. 6c Lack, E. (eds.) 1985. A Dictionary of absence, were likewise fundamentally a matter of Birds. Calton, UK:T. &A. D. Poyser. & scrutinising the evidence (see Collar Stuart Collar, N. J. 6>C Andrew, P. 1988. Birds to Watch: The 1985) and patiently covering the ground in the ICBP World List ofThreatened Birds. Cambridge, & most likely places (Ash Gullick 1990, Sergeant UK: International Council for Bird Preservation. etal. 1992). These three rediscoveries all occurred Collar, N. J. 6c Stuart, S. N. 1985. ThreatenedBirdsof after considerable prior ornithological activity in Africa and Related Islands: the ICBPHUCN Red the vicinity of the respective type localities, and DataBook. Cambridge, UK: International Council theyspeakto uswith thesame single message: it is for Bird Preservation 6c International Union for still too soon to place Pogoniulus makawai in syn- Conservation ofNature andNatural Resources. onymywith P. bilineatus and ifwe look long and Collar, N.J., Crosby, M. J. 6c Stattersfield,A.J. 1994. , hard enough we mayyet be pleasantlysurprised. Birdsto Watch2: TheWorldListofThreatenedBirds. Cambridge, UK: BirdLife International. Acknowledgements Collar, N. J. 6c Rudyanto 2003. The archive and the MarkAdamsoftheNatural HistoryMuseum,Tring, ark: bird specimen data in conservation status UK, answeredseveralqueriesfollowingourexamina- assessment. Bull Br. Omithol. Cl 123A: 95-113. tion ofthe type specimen ofP. makawai to which Crowe, T. M., Keith, S. 6c Brown, L. H. 1986. Robert Prys-Jones allowed us access,. Claire Phasianidae, guineafowl, Congo Peacock, quail, Spottiswoode commented on several drafts of this partridges and francolins. In Urban, E. K., Fry, C. paper, took two ofthe photographs, and suggested H. 6c Keith, S. (eds.) The Birds ofAfrica. Vol. 2. the comment concerning melanism, whilst Nick London, UK:AcademicPress. & Mundy checked and approved that comment. Dickinson, E. C. (ed.) 2003. The Howard Moore Subsequent drafts were read and commented on by Complete Checklist ofthe Birds ofthe World. Third John Colebrook-Robjent, Tim Dodman, Bob edn. London, UK: ChristopherHelm. Dowsett, Fran^oise Dowsett-Lemaire, Pete Leonard, Dowsett, R. J. 1973. New distributional records from TerryOatleyand Michael Stuart Irwin. We are most North-western Province. Bull. Zambian Omithol. grateful for all this help. Soc. 5: 66-69. Dowsett; R. 1980. Comments on some ornithologi- J. References cal type-localities in Zambia. Zambia Mus. J. 5: Ash,J. S. 6cGullick,T. M. 1990.Serinusflavigularedis- 7-16. covered. Bull. Hr. Omithol. Cl.\10: 81-83. Dowsett, R. J. 6c Dowsett-Lemaire, F. 1980. The sys- Aspinwall, I). 6c Bed, C. 1998. A Field Guide to tematic status ofsome Zambian birds. Gerfaut70: ZambianBirdsnotfoundinSouthernAfrica. Lusaka: 151-199. Zambian Ornithological Society. & 26-BullABCVol13No 1(2006) WhatisPogoniulusmakawai?: Collar Fishpool Dowsett, R. J. & Dowsett-Lemaire, R 1993. Oatley,T. B. 1969. Birdecologyintheevergreenforests Comments on the taxonomy ofsome Afrotropical ofnorthwestern Zambia. Puku 5: 141-180. birdspecies&. TauracoRes. Rep. 5: 323-389. Ryan, P. & Ca—ssidy, R.&2003.The ghost in the mavun- Dowsett, R.J. Forbes-Watson,A. D. 1993. Checklist da.Africa Birds Birdingl(6): 37-41. ofBirdsoftheAfrotropicalandMalagasyRegions, 1; Sergeant, D. E., Gullick,T.,Turner, D.A. &Sinclair,J. Species Limits and Distribution. Liege: Tauraco C. 1992. The rediscovery of the Sao Tome Press. Grosbeak Neospiza concolor in south-western Sao Dowsett-Lemaire, F. 1988. Fruit choice and seed dis- Tome. BirdConserv. Intern. 2: 157-159. semination bybirds andmammals in the evergreen Short, L. L. & Horne, J. F. M. 1985. Social behavior forests of upland Malawi. Rev. Ecol. (Terre et Vie) and systematics of African barbets (Aves: 43: 231-285. Capitonidae). Pp. 255-278 in Proc. Intern. Symp. Evans, M. I. 1991.The Red-tailedNewtoniaNewtonia Afr. Vertebr. Bonn: MuseumA. Koenig. fanovanae in the Ambatovaky Reserve, north-east Short, L. L. & Horne,J. F. M. 1988. Capitonidae, bar- & Madagascar. BirdConserv. Intern. 1: 47-52. bets and tinkerbirds. In Fry, C. H., Keith, S. Fjeldsa,J. 2003. PatternsofendemisminAfricanbirds: Urban, E. K. (eds.) The Birds ofAfrica. Vol. 3. how much does taxonomy matter? Ostrich 74: London, UK:AcademicPress. 30-38. Short, L. L. & Horne, J. F. M. 2001. Toucans, Barbets Ginn, P., Mcllleron, W. G. & Milstein, P. le S. 1989. andHoneyguides. Oxford: OxfordUniversityPress. The CompleteBook ofSouthernAfrican Birds. Cape Short, L. L. & Horne, J. F. M. 2002. Family Town: StruikWinchester. Capitonidae (barbets). In del Hoyo, J., Elliott, A. Goodman, S.M. &Schulenberg,T. S. 1991.Theredis- & Sargatal, J. (eds.) Handbook ofthe Birds ofthe covery of the Red-tailed Newtonia Newtonia World.Vol. 7. Barcelona: LynxEdicions. fanovanae in south-eastern Madagascar with notes Sibley, C. G. & Monroe, B. L. 1990. Distribution and on the natural historyofthe genus Newtonia. Bird Taxonomy ofBirds ofthe World. New Haven & Conserv. Intern. 1: 33-45. London, UK:Yale UniversityPress. Goodwin, D. 1965. Some remarks on the new barbet. Sinclair, I. & Ryan, P. 2003. BirdsofAfricaSouthofthe Bull. Br. Ornithol. Cl. 85: 9-10. Sahara. CapeTown: StruikPublishers. Irwin,M. P. S. 2003. ReviewofShort&Horne (2001). aConservation Biology Group, DepartmentofZoology, Honeyguide49: 237-238. LeonEavradn,s,P.MM.. I2.00(1ed.s.Z)aImmbpioar.taInntFiBsihrpdooAlr,eaLs.iDn.ACf.ric&a CCUonBui2rvte,r3sEiJGt,iyrtoUofKn,CaRamobnarddi,dBgiCerad,mLibDfroeiwdLngniteenrgCnaBStt3iroeneOatN,l,AC,aWemlUblrKbirdoEgo-ek andAssociatedIslands:PrioritySitesforConservation. Newbury: Pisces Publications & Cambridge, UK: mail: [email protected] bBirdLife Lnternational, Wellbrook Court, Girton BirdLife International. Leonard, P. M. 2005. ImportantBirdAreas inZambia. Road, Cambridge CB3 ONA, UK. E-mail: Lusaka: Zambian Ornithological Society. [email protected] Mayr, E. 1971. New species of birds described from 1956 to 1965./. Ornithol. 112: 302-316. Received 7June 2005; revision accepted 11 November Mundy, N. I. 2005. Awindow on the genetics ofevo- 2005 lution: MC1R and plumage colouration in birds. Proc. Roy. Soc. Lond. B272: 1633-1640. WhatisPogoniulusmakawai?: Collar&Fishpool BullABCVol13No1(2006)-27

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