<I>NEOPHRON PERCNOPTERUS</I>

j. RaptorRes. 36(1):17-23 ¸ 2002 The Raptor ResearchFoundation, Inc.

DESCRIPTION OF A NEW SUBSPECIES OF THE EGYPTIAN VULTURE (ACCIPITRIDAE: NEOPHRON PERCNOPTERUS) FROM THE CANARY ISLANDS

jOSl•ANTONIO DON•.ZAR, 1JUAN Josf• NEGRO, Cf•SAR JAVIER PALACIOS AND LAURA GANGOSO Departmentof AppliedBiology, EstaciSn Biol6gica de Do•ana, ConsejoSuperior de InvestigacionesCientificas, Avda M a Luisa sin 41013 Sevilla,Spain

JOSl•ANTONIO GODOY Laboratoryof MolecularEcology, Estaci6n Biol•gica de Do•ana, ConsejoSuperior de InvestigacionesCient(ficas, Avda M a Luisa sin 41013 Sevilla,Spain

OLGA CEBALLOS AND FERNANDO HIRALDO Departmentof AppliedBiology, Estaci6n Biol6gica de Dohana, ConsejoSuperior de Investigaciones Cientificas, Avda M • Luisa sin 41013 Sevilla,Spain

NIEVES CAPOTE Laboratoryof MolecularEcology, Estaci6n Bioldgica de Do•ana, ConsejoSuperior de InvestigacionesCient•cas, Avda M • Luisa s/n 41013 Sevilla,Spain

ABSTRACT.--Onthe basisof four studyskins from museumcollections and 37 live birds fkom the island of Fuerteventura,we describea new subspeciesof the EgyptianVulture (Neophronpercnopterus majorensis) from the Canary archipelago.Canarian Egyptian Vultures are significantlylarger than WesternEuropean and North Atkican individuals. In addition, some genetic differentiation may exist; analysesof the mi- tochondrialDNA control region revealedthat there are haplotypesexclusive to the CanaryIslands. The current population of EgyptianVultures in the Canary Islandsis 25-30 breeding pairs restrictedto the islandsof Fuerteventuraand Lanzarote,and this subspeciesis therefore endangered. KEYWOP, DS: EgyptianVulture; Canary Islands; control region; mitochondrial DNA; Neophron percnopterus majorensis;subspecies'.

Descripci6nde una nueva subespeciede alimoche (Neophronpercnopterus majorensis) nativa del archipi6- Iago de Canarias

RESUMEN.--Describimosuna nueva subespeciede alimoche (Neophronpercnopterus majorensis) nativa del archipi61agode Canarias,sobre la base de 4 pieles de museoy 37 individuoscapturados vivos. Los alimochescanarios son significativamente mayores que los del oeste de Europay nortede •f?ica. Ade- mas, presentan diferenciaci6n gen6tica:un analisisde la Regi6n Control del ADN mitocondrial mostr0 que existen haplotipos exclusivosde las Islas Canarias.La poblaci6n actual de alimochescanarios es de 25-30 parejasrestringidas alas islasde Fuerteventuray Lanzarote,y estfi,por tanto,gravemente amenazada. [Traducci0n de los autores]

The genusNeophron has only a singlespecies, the curring in most of the range of the species:Eu- Egyptian Vulture (Neophronpercnopterus), which rope, Africa, Middle East, central Asia, and lives in dry ecosystemsof the Palearctic, Ethiopic, northwestIndia, and N. p. ginginianusin most of and Indo-Malayanbiogeographic regions. Two sub- the Indian subcontinent (Brown and Amadon specieshave been recognized:N. p. percnopterusoc- 1968, del Hoyo et al. 1994). Following Brown and Amadon (1968) the latter is slightlysmaller, having yellow as opposed to dark-brown bill and compar- E-mail address: [email protected] atively-weakerfeet and claws.

17 18 DON•ZAR ET AL. VOL. 36, NO. 1

The Canary Islands are an archipelago of vol- of Natural History (BMNH) collection (Tring Cat- canic origin formed in the past 20 million years alogue No. 199). close to the western Afi:ican coast (100 km for Geographical Distribution. The subspeciesis Fuerteventura, the nearest island). The islands currently endemic to the islands of Fuerteventura have a subtropicalclimate; fauna and flora are re- and Lanzarote (eastern Canary Islands). It almost lated to other Macaronesian archipelagos (Madei- certainly occurred in the remaining islands of the ra, Azores) and to the Mediterranean region. archipelagountil extirpated in the 20th century. Twenty-sevenpercent of the vascularplants and There are three specimensin the British Museum 50% of the invertebratesare endemic (Juan et al. of Natural History (Tring) which were collected at 2000). In addition, sevenavian speciesare consid- Gran Canaria (BMNH No. 121) and Tenerife ered to be endemic of the archipelago (Blanco and (BMNH No. 231 and BMNH No. 232, respectively) Gonz•dez 1992). Endemic subspecieshave already at the beginning of the 20th century. been describedfor three birds of prey living in the Description. Plumage patterns of color of Neo- islands: Common Buzzard (Buteo buteo insularum, phron percnopterusmajorensis resembles the nomi- Foericke 1903), Eurasian Sparrowhawk (Accipiter nate subspeciesN. p. percnopterus(Brown and Area- nzsusgranti, Sharpe 1890), and Eurasian Kestrel don 1968, Cramp and Simmons 1980). Adult (Falco tinnunculus canariensis,in the West islands, individuals of the Canarian subspeciestypically Koenig 1890; and E t. dacotiaein the East islands, show white plumage impregnated by rufous col- Harterr 1913). Three other raptor speciesexist in oration, especially in the crown, nape, median the archipelagowithout known distinct subspecific wing coverts,breast, and tail. This colorationis var- status: Egyptian Vulture, Osprey (Pandion haliae- iable among individualsand seemsto be acquired tus), and Barbary Falcon (Falcopelegrinoides) (del from iron oxides derived from the local soils, Hoyo et al. 1994). which are rich in iron compounds;evidently, this In Macaronesia (Palearctic subregion including is an ecologicalattribute associatedwith this pop- the Atlantic Islands), the EgyptianVulture inhabits ulation and not a taxonomic characteristic. Can- the Canary and Cape Verde archipelagos(Banner- arian Egyptian Vultures are sedentary and thus man 1963, Bannerman and Bannerman 1965, differ behaviorallyfrom Western European popu- 1968). In the Canary Islandsthis bird wascommon lations, which are long-distancemigrants (Cramp historically;the speciesformerly occurred in the and Simmons 1980). •slands of La Gomera, Tenerife, Gran Canaria, Measurements.We captured and measured37 Fuerteventura and Lanzarote (Martin 1987). At wild individuals in 1999-2000 (all birds released present, the Egyptian Vulture persistsonly in the two easternmost islands (Fuerteventura and Lan- after capture), and compared these data from individualsfrom continental Spain (Table 1). Mea- zarote) with a total population estimatedat 25-30 pairs in the year 2000 (pets. observ.). surements (mm) of the holotype (BMNH-Tring We have observedstrong differences in the mor- 199) are: flattened wing chord 516.3 mm; tarsus phology (see below) of this population in relation 88.2 mm; tail: 240.6 mm; bill length with cere 63.5 to that of Western Europe and North Africa. In mm; bill length to the distal edge of cere:31.8 mm. addition, analysesof the mitochondrial DNA con- Morphological Comparisons.N. p. majorensisdif- trol region revealed that haplotypesexist exclusive fers from the nominate subspecieson the basisof to the Canary Islands.This result suggeststhat the larger body measurements.Comparisons with live population has been isolatedfrom othersfor a very Iberian individuals (Table 1) revealed significant long time. Consequently,we propose the recogni- differencesfor every trait when the effectsof age tion of this population at the subspecificlevel. In and sex were controlled. Differenceswere partic- this paper, we describethe Canarian EgyptianVul- ularly large for body mass,with the island birds tures as Neophronpercnopterus majorensis subsp. nov. being 18% heavier than Iberian vultures. Tail feathersand wing chord were about 4-8% longer DESCRIPTION in Canarianvultures. Smaller (ca. 2%), but still sig- Holotype. This specimenis an immature (sec- nificant differenceswere detected in the length of ond plumage) male collectedon 22 October 1913 the tarsus,primary-feather, bill, and bill with cere. in Tost6n (northwestern coastof Fuerteventura) by Gigantism is well known to occur on islands (Pe- D. Bannerman. It is located in the British Museum tren and Case 1997, Grant 1998), which has been MARCH 2002 SUBSPECIESOF EGYPTIAN VULTURE 19

attributed to the absence of dominant species (Thaler 1973). We lack measurements of birds from neighboring "Sahara," where the small residentpopulauon seems to have been extirpated (see below). We have examined one live bird from the former SpanishSahara (now Morocco),captured in 1975, which is currently at the Zoo of Jerez (Spain). This bird's measurements (mm) were: wing chord = 505, tail = 256, tarsus= 75.9, primary = 376, bill length = 31.1, bill cere = 59.4, and mass = 1820 g. These measurementsare similar to those of continental birds from the Iberian Peninsula (Table 1). Measurements from existing museum specimens are not reliable for the description of the Canarian subspeciesas there is only a single skin from Fuer- teventura and the taxonomic status of the extirpated EgyptianVultures from other GanaryIslands is unknown. In addition, problems relative to the condition of skins, small sample size, and the ex- istenceof variability linked to age and sex preclud- ed statisticalanalyses. Nonetheless, we took measures of flattened wing chord (the measure that can be taken with least error in our experience) from specimensdeposited at the BMNH (Tring) (Table 2). The four Ganarian Egyptian Vulture skinsshowed the highest valuesfor wing-chordfor the entire range. Their values were well above those of individualsfrom neighboring populations in the western Mediterranean, continental Africa, and Cape Verde Islands. This suggeststhat birds from the Canary Islandsconstituted a single subspecies,which is today restricted to Fuerteventura and Lanzarote. There are no additional skins de- positedin other museums,which precludesfurther comparisons.It seemsinteresting that Cape Verde EgyptianVultures are smallerthan Canarian birds. Hazevoet (1995) reported that these birds are similar in plumage to individuals from Afi:ican populations; he considered them as belonging to the nominal subspecies.

ET1MOLOGY Scientific Name. The name of the proposed new subspeciesis derived from "Majorata," the ancient name of Fuerteventura Island. It was so called by the Spanish conquerors, since the main native guanche tribe of the islandwas known as the "Ma- jos." At present, the inhabitants of the island are still called "majoreros." English Name. According to the species geo- 20 DONXZAR ET AL. VOL. 36, No. 1

Table 2. Flattened wing chord measurements (mm) of Egyptian Vulture (Neophronpercnopterus) skins kept in the British Museum of Natural History (Tring).

REGION SUBSPECIES MEDIAN RANCE N

CanaryIslands N.p. majorensis 505.7 490.0-516.3 4 Cape Verde Islands N.p. percnopterus 489.6 456.0-515.6 6 WesternMediterranean a N.p. percnopterus 490.0 472.0-510.0 14 EastAfrica b N.p. percnopterus 480.4 450.0-518.0 13 M•ddle Eastc N.p. percnopterus 501.3 485.5-505.0 4 SouthArabia d N.p. percnopterus 475.7 351.1-491.3 4 Indian subcontinent N.p. percnopterus 493.5 471.0-510.0 14 Indian subcontinent N.p. ginginianus 460.6 441.5-495.3 23 Spain, France,Algeria, Morocco. Abyssinia,Somalia, Kenya. Egypt,Palestine, Turkey. Oman, Socotra, Arabia. graphical distribution of this subspecieswe pro- heritance of the analyzed sequences,indicating pose the name Canarian EgyptianVulture. that these were of mitochondrial origin and not from nuclear insertions. gENETIC DIFFERENTIATION Results.A fragment of 459 base pairs of control Methods. We used sequencedata of the control region wasused in the sequenceanalyses. Sequenc- region of mitochondrial DNA from 11 Egyptian es for the 11 individuals were all different, i.e., Vultures of different provinces (two from Fuerte- each individualpresented a unique haplotype(Fig. ventura, two from the Balearics, four from conti- 1). A total of 53 polymorphic siteswere observed nental Spain, one from westernSahara [Morocco] (11.5% of the 459 bp sequenced).Absolute num- and two from India [N. p. ginginianus](Table 3). bers of pair-wisedifferences ranged from 2 (0.44%) Analysesof mtDNA have been particularly com- between samples from Canary Island and 35 mon for studiesof subspeciesand closelyrelated (7.63%) between Iberian and Indian samples.Net speciesbecause of the relativelyrapid rate of evo- mean Kimura's 2 parameter distancesbetween In- lution and ease of analysis relative to nuclear dian and other groupsranged from 0.066 to 0.077, (chromosomal) DNA (Wilson et al. 1985, Avise et while the range of distancesbetween non-Indian al. 1987). groups was 0.005 (Iberian and Balearic) to 0.020 Total DNA wasextracted from blood samplesfol- (Canarian and Saharian). Consequently,phyloge- lowing Gemmell and Akiyama (1996) with some netic analysesof these sequenceswith using differ- modifications.Preliminary work in our laboratory indicated that the order of genesin the mitochon- ent algorithms approaches (maximum parsimony, minimum evolution, and maximum likelihood) drial molecule of Egyptian vultures is consistent with the new avian gene order recently described consistentlygrouped the two Indian sequencesin by Mindell et al. (1998). We thus targeted for am- a clade widely separatedfrom the rest of Iberian, phfication by PCR a potentially hypervariablefrag- Balearic, Canarian or Saharian sequences,with ment located between the conserved sequence bootstrapvalues close to 100% (Fig. 1). The two block called Fbox and the flanking Thr-tRNA (for Canarian sequencesalso consistentlygrouped to- primer sequencesand PCR conditions please con- gether as a subcladewithin non-Indian sequences, tact authors). Sequencingreactions were run in an with bootstrapvalues up to 92%. A clade of Bale- ABI-377 automatic sequencingsystem (Applied aric sequenceswas obtained with some algorithms, Biosystems)in an external laboratory. Sequences with valuesof up to not higher than 62%. Finally, were manually edited and aligned. Genetic dis- the only Saharian sequence included was more tance calculationsand phylogeneticanalyses were closelyrelated to Iberian or Balearic than to Can- performed with the program PAUP-Version arian sequences.The sequencesare deposited at 4.Ob4a (Swofford 2000). Previousanalysis of fam- the EMBL databank under accession numbers ilies of captive birds showeda strict maternal in- AJ305147to AJ305150(N. p. majorensis),AJ305151 M_ARCH2002 SUBSPECIESOF EGYPTIANVUI,TURE 21

Canary1

Canary2

Ibena 1

Ibena 3

Iloena 2

E•.learlc 1

E•leanc 2

Sahara 1

Iberia 4

Ind•a 1

1133

, India 2 0005 changes Figure1. Phylogeneticrelationships among the Egyptian Vulture Control Region sequences identified in thisstudy Minimumevolution tree, constructedwith the programPAUP using Kimura's 2 parameterdistances and the neighbor-joiningalgorithm followed by a branch-swappingprocedure. Values above branches represent support from 100 bootstrapreplications (only values above 50% are shown). to AJ305162(N. p. percnopterus),and AJ305163to percnopteruson the basisof its largersize (Table 1, AJ305166 (N. p. ginginianus). 2); the difference is strong with respect to Iberian EgyptianVultures, which constitutethe main west- DIAGNOSIS ern Palearcticpopulation. These latter birds mi- The new subspeciescan be distinguishedfrom grate along the African coast (Cramp and Sim- individualsbelonging to the nominal subspecies mons 1980) and, thus, have some chance of 22 DONAZA• ET AL. VOL. 36, No. 1 straggling to the Canary Islands. Morphological show them closer to continental birds. This would differences between majorensisand percnopterusof not invalidate, however; the existence of a differ- western European and African populations (in- entiated Canarian lineage as it is shown in this pa- cluding CapeVerde Islands)are asmarked asthose per. existing between the subspeciesginginianus and CONSERVATION percnopterusin central Asia (Table 2). A wider population genetic screeningof the spe- As was stated above, the entire Canarian Egyp- cies will be needed to assesslevels of genetic vari- tian Vulture population is restricted to Fuerteven- ability and estimationsof gene flow among sub- tufa and Lanzarote, where no more than 30 terri- populations. Meanwhile, the analysisof Control tories remain occupied. Total population has been Region sequencesin a limited number of Egyptian estimated at around 130 birds (pets. observ.). Vultures showslimited low genetic diversitywithin Breeding successhas been extremely low in recent the Canary Islands populations versus the larger years (ca. 0.5 fledglings/pair in 1998-9000). Cur- Iberian populations. On the other hand, the Can- rent threats to this population include mortality arian population showsunique mitochondrialhap- from power lines due to collision and electrocution lotypes that group monophyletically within N. p. (12 casesduring the studyperiod; see alsoLorenzo percnopterusnon-Indian sequences.Both resultsare 1995), poisoning (4 cases), and lead intoxication consistentwith colonization and further expansion by ingestion of lead bullets (pets. observ.). Theft in the islands by individuals from the mainland of eggsand young at the nest, and other human and suggestlimited gene flow between Canarian activitiesmay account for some casesof nest de- and other populations. A differentiated evolution sertion during the breeding period (Palacios of the Canarian population of Egyptian Vultures 9000). Conservation measures should be directed would have been favored by the isolationresulting preferentiallyto prevent casualtiesrelated to power from the 100 Mn-wide sea corridor existing be- lines, lead intoxication, and illegal poisoning. Re- tween the eastern coast of Fuerteventura and that inforcement with individuals from continental ar- of the African continent. Possibilityof individual eas is not recommended, on the basis of the ge- interchange with other breeding populations ap- netic differences showed by the Canarian birds pears to be low. The current population in the with respect to those of neighboring populations. western Sahara seems to be virtually extinct (J. ACKNOWLEDGMENTS Don•tzar unpubl. data), and therefore the main possibility of genetic exchange would seem to The Consejeria de Medio Ambiente del Cabildo Insu- come from Iberian birds migrating along the Af- lar de Fuerteventura and the Project PEN 2000-1556 GLO funded this research.Our specialthanks to Manuel rican coastand stragglingto the islands.We cannot Miranda and Carlos Alba who stimulated the project and •ule out the possibilitythat some Iberian individ- provided logistic support. Gorgonio Diaz and Maria uals reach the island and eventually interbreed Asunci6n Delgado (Consejeria de Politica Ambiental, with local individuals. However; there is strong ev- Gobierno de Canarias) gave scientific banding permits We thank very much Manuel de la Riva, Jos6 A. S•tnchez idence that Fuerteventura'sEgyptian Vultures con- Zapata, Gema Garcia, Reyes L6pez-Alonso, Manuel V•tz- StltUtean ecologically-isolatedpopulation with well- quez, Yohama Enriquez, Ana Calero, Gema Mosquera, differentiated morphological and genetic Nemesio Hern•mdez, D'•maso Santana, Francisco Garcia, characteristics. Jos6 Hern•mdez, and Eduardo Castilla for assistance Summarizing, we propose that the naming of throughout the research.DNA sequencingas well as the sexing of live-trapped birds was done in the Laboratory this new subspeciesis justified. Description of new of Molecular Ecology (Estaci6n Bio16gicade Dofiana). subspecies,and even of avian species,has been re- Drs. Mark Adams and Robert PrysJonesfrom the British cently done on the basis of only biometric ap- Museum of Natural History (Tring) and Manuel Barcell proaches and taking in account only a few speci- (Zoo of Jerez de la Frontera) permitted accessto specimens in their care. Dean Amadon,Jos6 L. Tella, and Clay- mens (e.g., Smith et al. 1991, Forero and Tella ton M. White provided valuable criticism of the manu- 1997, Safford et al. 1995, Preleuthner and Gamauf script. 1998, Yosef et al. 9000). In some casesthe distance between these new taxa and those closelyrelated LITERATURE CITED were lower than that found in our study.We cannot AVISE,J.C.,J. ARNOt.D, R.M. BALL,E. BERM1NGHAM,T. LAMB, discard that further genetic analysesmight permit J.E. NEIGEL,C.A. REEB,AND N.C. SAUNDERS.1987. In- the discoveryof additional haplotypeswhich may traspecific phylogeography:the mitochondrial DNA MAP,C}• 2002 SUBSPECIESOF EGYPTIAN VULTURE 23

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