© Biodiversity Heritage Library, http://www.biodiversitylibrary.org/; www.zoologicalbulletin.de; www.biologiezentrum.at Bonner zoologische Beiträge Band 51 (2002) Heft 2/3 Seiten 97-104 Bonn, September 2003 Museum Collections as Sources of Genetic Data & Robert B. Payne Michael D. Sorenson Museum ofZoology and Depailment ofEcology and Evolutionary Biology, University ofMichigan, Ann Arbor & Department ofBiology, Boston University Abstract. Museum collections are an under-utilized source ofgenetic material for avian systematics. Museum specimens are particularly valuable when the collection ofnew material is difficult or impossible. This is the case for rare or extinct birds and forregions wherecollecting is notallowed. Museum collections alsoprovide opportunities forthe studyofrecent evolutionary change. First, we illustrate the importance ofvoucher specimens for tissue samples collected for molecular systematics studies, reportingon several cases inwhicherrors intheoriginal identificationwerecorrectedaftergeneticdata prompted a re-examination ofthe voucher specimen. Second, we report results from our molecular systematics studies of finches Viduidae and Estrildidae and cuckoos Cuculidae in which older museum specimens provided the genetic samples for many taxa. Finally, we discuss the use offeathers taken from museum specimens in molecular studies. Working with older feathers rather than fresh tissues entails additional work in the laboratory but has advantages in identification and repeatability, and museum skins are often the only material available for less-common species. The length ofPCR product thatcan be amplified from featherextracts declines with specimen age, such that genes mustoften be sequenced in smaller segments (e.g., 200-300 base pairs, bp) in specimens that are 30 to 50 years old. We have consistently obtained mitochon- drial DNA sequence data from specimens dating back to 1950 and have had success with a limited numberofolder speci- mens. Although the probability ofsuccess in amplifying a target gene declines with age, some skins in any age class may yield usable products for sequencing (the "universal primers" that often are used forcytochrome-b may be less useful than specific primers). Our success in amplifying and sequencing genes from museum skins encourages the use ofcollections for sampling critical specimens. The management ofscientific resources includes both archival safekeeping ofspecimens collected from the field and the considered use ofspecimens in systematic research. We suggest that the association of genetic data gives specimens "value added" status and urge research museums to consider the benefits in addition to the costs ofusing specimens as primary sources ofcomparative genetic material. Key words, ornithological collections, DNA, phylogeny, voucher specimens 1. DNA FROM MUSEUM SPECIMENS Using bird skins in museums as a primary source of genetic information has a number ofadditional advan- Both archival and scientific use require the wise allo- tages. Skins are a potential source ofgenetic samples cation ofvaluable museum specimens. After an initial enthusiasm for recovery ofDNA from museum speci- for birds that are not possible to collect in the field, mens (HouDE & Braun 1988), museum curators real- due to extinctions or political considerations. Museum specimens also can be used to test repeata- ized there is a potential conflict between archival main- bility (a second sample can be taken from a known tenance and progressive management of a scientific resource (Graves & Braun 1992). With their term single source), can be examined to verify identifica- tion (Ruedas et al. 2000) and to provide new infor- „destructive sampling" for the use ofskin and feathers from museum specimens, Graves & Braun (1992) mation as molecular techniques develop and new bio- logical questions arise. demand the researcher tojustify the need for sampling an irreplaceable specimen. At the same time they rec- Ofcourse, material that is collected in the field specif- ognize their importance for species that otherwise are ically for genetic analysis is generally ofhigher qual- unavailable for research. In response to increased inter- ity and utility than material that can be recovered only est in sampling specimens for genetic research, many with much labor from museum specimens (Arc- research museums have established guidelines for sam- TANDER & Fjeldsá 1994). Tissue samples for genetic pling and criteria for museum curators to evaluate these analysis should always be saved when a bird is col- proposals (USNM, FMNH, UMMZ). An alternate tenn lected and a specimen is prepared. Muscle tissue or for the activity is „value added sampling" because the blood can be preserved in liquid nitrogen (LN2) or in information acquired about a specimen's genotype con- suitable buffer (Seutin et al. 1991), or even in etha- tributes to the growth of scientific knowledge and the nol. A well-designed research program includes pre- potential to solve biological problems. The specimens served samples ofboth blood and muscle tissue and a gain invalueto ourscientific community when they are specimen, both as a voucher for identification pur- used in this manner poses and for its value in augmenting existing natural © Biodiversity Heritage Library, http://www.biodiversitylibrary.org/; www.zoologicalbulletin.de; www.biologiezentrum.at 98 Robert B. Payne & Michael D. Sorensen history collections apart from its immediate applica- SON et al. 1999; Cracraft & Feinstein 2000; Dumb- tion to a particular genetic study. The cost in money acher & Fleischer 2001; Sefc et al. 2001, 2002; and time required to obtain permits for scientific work Payne et al. 2002). Recently collected skins are more is about the same for trapping birds and collecting useful than older skins (Glenn et al. 1999). Our only genetic samples as it is for collecting complete research with finches and cuckoos has had high suc- specimens. As the scientific value is greater for the cess in recovering genetic sequence data from speci- complete sample, we encourage molecular systema- mens collected back to 1900, but success in recover- tists to secure their own genetic samples in the field ing data from the earlier specimens (e.g, < 1960) and to collect new museum specimens for systematic required amplification of relatively short PCR frag- research (Remsen 1995). ments (e.g., < 200-300 base pairs in length). In other laboratories, genetic information has been recovered Nevertheless it is not always possible to get fresh from bird skins collected as long ago as 1860 and material for genetic studies. Moreover, certain evolu- tionary questions involve sampling museum speci- 1874 (Ellegren 1992, 1994; Prinzinger et al. 1997). mens to test evolutionary change through time (e.g., Birds collected in recent years are often taken specif- Thomas et al. 1990; Glenn et al. 1999) and to iden- ically for preserved tissues to be used in genetic stud- tify historically important material such as type spec- ies. In many studies, voucher specimens (skins or imens (Prinzinger et al. 1997). Little work has been skeletons, or both) are retained to support the identifi- DNA done with older specimens, because degrades cation; a multiple preparation is most useful (Winker with age or because the skins were preserved with 2000). Specimens preserved in alcohol can be vouch- chemicals such as arsenic (Pääbo 1990; Ellegren ers for later identification, but specimens preserved in 1994). The past decade has made increasing use of PCR amplification to obtain sequences from minute formalin are less useful for genetic analyses because of the difficulty of extracting and amplifying DNA. fossil or forensic amounts. The quality of preserved material necessary to do this work is not nearly as Recovery of genetic sequence is more difficult when restrictive as the liquid nitrogen-preserved tissues skeletons are cleaned of all connective tissue and generally used in restriction site analysis (Hillis et al. muscle. Retaining some connective tissue on a skele- 1996; Klein & Payne 1998). ton makes the specimen more useful both for anatom- DNA ical research and genetic analysis. can be extracted, amplified and sequenced from museum study skins and skeletons both for phyloge- For molecular genetics work we prefer to use muscle netic analysis and for population-level work with tissue that was preserved in the field. When fresh tis- microsatellites (e.g. Taberlet & Bouvet 1991; Elle- sue is not available, we prefer feather samples over gren 1992; Leeton et al. 1993; Cooper 1994; blood to avoid the risk of sampling nuclear copies of Cooper et al. 1996, 2001; Mundy et al. 1997; mitochondrial genes (Sorenson & Quinn 1998). Prinzinger et al. 1997; Engstrom et al. 1999; Soren- Nuclear copies of mtDNA sequences (or 'numts'; Lopez et al. 1994) result from ancient transpositions of mitochondrial sequences into the nuclear genome. Tab. 1: Table ofsuccess (or sequence length recovered) vs age of Over time, the mitochondrial and numt sequences specimen diverge and evolve at the different rates ofmolecular Age of # samples #samples #yielding PCR evolution characteristic ofthe two genomes. In some specimens extracted sequenced products <500 hp cases, a numt resulting from an ancient transposition in length event is present in a group of related species des- Cuckoos cended from a common ancestor (e.g., Arctander 1930 - 1939 1 1 0 1995; Bensasson et al. 2001), while in others multi- 1940 - 1949 2 2 1 ple transposition events have occurred within a single 1950 - 1959 16 14 8 & clade (Sorenson Fleischer 1996). In either case, 1960 - 1969 30 26 17 1970 - 1979 6 6 6 numts are a potential source ofconfusion in compar- 1980 - 1989 11 9 8 ative analysis of mtDNA, the genome that has been 1990 -2001 18 18 18 used most widely in avian molecular systematics. The Estrildidae high ratio of mtDNA to nuclear DNA in muscle and 1950 - 1959 5 5 5 feathers makes these tissues preferable to blood when & 1960 - 1969 15 13 13 mitochondrial genes are of interest (Sorenson 1970 - 1979 9 9 9 Quinn 1998). Feathers are a good source of mtDNA. 1980 - 1989 6 4 4 1990 - 2001 45 44 44 A feather sampled from a museum skin can be © Biodiversity Heritage Library, http://www.biodiversitylibrary.org/; www.zoologicalbulletin.de; www.biologiezentrum.at Museum Collections as Sources ofGenetic Data 99 replaced after the base ofthe quill is sampled, much Anomalospiza as feathers dropped in the preparation ofa study skin. The brood-parasitic cuckoo-finch Anomalospiza im- Careful selection of a feather or feathers to sample berbis was once thought to be a ploceid finch, as its (e.g., a contour feather from the back ofthe bird or an plumage is like that of weavers Ploceus and bishops underwing covert or axillaiy) results in little or no Euplectes (Sibley & Monroe 1990). Its relationships effect on the specimen's appearance and negligible are of interest because Anomalospiza and the indigo- damage to the specimen. Because toepad morphology birds and whydahs Vidua are the only Old World is potentially infoimative about avian relationships brood-parasitic songbirds. Understanding the rela- and adaptation (Clark 1973), removing a toepad for tionships of Anomalospiza, Vidua, and their nesting genetic analysis is not necessarily less destructive to a relatives is needed to test ideas about the evolution of specimen than removing a feather. brood parasitism (Payne 1998). Anomalospiza feath- UMMZ Costs for the museum collection as a primary source ers were sampled from museum skins col- ofgenetic material are twofold: lected in 1968 and 1972 and processed in standard 1. Damage to specimens. Museum curators can extraction, amplification and sequencing methods. Nucleotide sequences were compared with those of determine whether specimens that are sampled ploceid, estrildid and viduid finches and with other will have significant value added, or whether the cost in terms ofdamage or even loss ofspecimens songbirds. Anomalospiza is most closely related to Vidua, and these two genera ofbrood-parasitic finches is greater than the expected scientific interest. form a lineage that is sister to the estrildid finches 2. Time and effort. Museum curators can negotiate Estrildidae, the group that includes the host species of whether the museum or the geneticist is responsi- Vidua. Brood parasitism evolved only once in the Old ble for removal of the feather, snip of skin, or World songbirds, and the common ancestor ofAnom- toepad. The museum curator may be more con- alospiza and Vidua dates to perhaps as long as 20 mil- servative in selecting the sample, whereas the lion years ago (Sorenson & Payne 2001). geneticist may be interested in obtaining suffi- cient material to carry out the molecular work with success. Taking very small samples is false Estrildidae economy ifthe sample yields no genetic data as a The estrildid finches Estrildidae comprise about 140 result. species. Biological questions in the group include reconstruction of their biogeographic history with GENETIC ANALYSIS OF MUSEUM 2. multiple dispersal events between Australasia and SPECIMENS RESOLVES PHYLOGENETIC Africa, and relationships among the estrildid genera QUESTIONS that are parasitized by Vidua finches, brood parasites Our results in the analysis ofmtDNA in feathers from that have frequently colonized new host species & museum specimens include several that are ofinterest within an estrildid genus (Klein Payne 1998). In addifion, the relationships ofspecies within the Estril- to avian systematics and emphasize the importance of didae are not well known. Specimens were obtained skins as primary sources of genetic information. by collecting tissues and feathers in the field, and by Results include the following: African Pholidornis sampling feathers from museum specimens and live nishiae is more closely related to the African warblers birds. For 33 ofthese estrildid species the source was than to the penduline tits or the estrildids (Sefc et a feather from a museum specimen (FMNH, MNHN, al.2002); a hybrid Vidua (Payne 1980) indigobird x UMMZ, ZMFK, ZMUC). paradise whydah had an indigobird as the maternal & parent (Payne Sorenson, in prep.); drongo cuckoos Surniculus are related to Cuculus rather than to koels Paludipasser locust finch Eudynamys; New Guinea white-crowned black A cuckoo Caliechthrus is closely related to Cacomantis feather resolved the evolutionary status of African and not to koels Eudynamys; and long-billed cuckoo locust finch Paludipasser locustella. The bird is basal Rhamphomantis longirostris is a Chrysococcyx to all other estrildid finches and it is not a waxbill {Sorenson & Payne, in prep.). Other results are men- (Estrildini). It was first described as a distinct genus tioned in more detail: Paludipasser by Naeve 1909, then was placed with © Biodiversity Heritage Library, http://www.biodiversitylibrary.org/; www.zoologicalbulletin.de; www.biologiezentrum.at 100 Robert B. Payne & Michael D. Sorensen & quail-finch Ortygospiza by Lynes Sclater (1934). chestnut-breasted mannikin Lonchura castaneotho- Locust finch resembles quail-finch Ort}>gospiza atri- rax). Paludipasser is not closely related to quail- collis in having barred flanks in the female, and a red finch, to other waxbills, or to another group ofestril- bill. Chapín (1954) noted, „in recent years these have did finches. It is an estrildid finch with no close rela- usually been treated as members of a single genus, tionship to other finches (Sorenson & Payne 2001b.). Ortygospiza, a course here followed with some reluc- tance... The beak ofP. locustella is much deeper, with Lonchura munias culmen more strongly ridged, and it actually shows a certain resemblance to that of Anomalospiza. The Results with the munia Lonchura species are incom- wings oílocustella seem to me much smaller, propor- plete, but preliminai7 results show the validity of tionally, and its powers of flight not nearly so great. genetic sequences recovered from older museum The legs, on the other hand, are exceptionally stout specimens and the hybrid origin of a recently and muscular in locustella, more so than in any other described bird, the cream-bellied munia Lonchura estrildine finch I have dissected. The form ofthe gape pallidiventer. The phylogenetic estimates ofrelation- wattles and the palatal spotting are still unknown, so 1 ships within Lonchura are consistent with those of suspect we have more to learn ofthe relationships of Baptista et al. (1999). Goodwin (1982), Réstale locustella.'''' (1997) and Baptista et al. (1999) all recommended The palate of nestling locust finch Paludipasser has including „Padda" oryzivora and „P. "fuscata in the an arc-shaped black line and a pair ofshort black lines genus Lonchura, consistent with our results that also behind the arc and at an angle to it. The gape has two include L. leucosticta and L. tristissima in this clade flat red lobes and the lining ofthe mouth is bright red (Fig. 1). (Irwin 1958). A study skin ofajuve- nile (FMNH 283598) was examined I coitamolirorea by softening the head in water. The L qnntxohr melanin palate marks and gape lobes L nevermaynt were visible as described by Irwin L ßav}prymna £.5p«r/afcí7its(¡977) (1958) though lacking the red colors. L ¡pictabjlii The palate of most waxbills has ^ L teermk spots, whereas the palate of grass- L grendii pallida finches (Poephilini) and munias and moja mannikins (Lonchurini) has lines J LL ppdalnidvctier (IMMELMANN et al. 1965,1977; 66 If 'i /- aricapiUa (1950) Réstale 1997). In contrast to locust M ' I / at tlnch, nestling quail-flnch Ort\>- *- LL mmddaacca nr- LL f/umicccm gospiza has six palate spots, the m. 95 P- I iiffrrtuata palate is whitish, and the gape has 100 J" Lr I / llee.uco^aitra three pale blue globes separated by '-• •£./. monutUi\967) 74 I CQ black in a checkerboard pattern. The L puyvnáata palate in locust finch is unlike the 90 J L /uxata waxbills and is more like the munias L oorryyTn3v\ora 100 latcoihcta and mannikins. The plumage pattern 94 tnstissma ofbarred flanks in locust finch occurs 100 Eu>dxe cantam Buodice malabanca in several estrildid finches, not only Ltrnvreahti »aia in waxbills (quail-finch Ortygospiza ICO *SptrmcstiiíSfriagütoida(1961 atricollis, green avadavat Amanclava 78 75 Spcrm^stei b hcolcr farinosa, goldbreast A. sub/lava) but 76 Sjxrmistei cixuUaTia also in Australian grassfinches Od>»toipaa ca»K<p¡ (plum-headed finch Aidemosyne 100 I He!<romi9»ap«:loral)i modesta, juvenile diamond flretail * HcteromianapicToraliJ Stagonopleura guttata) and several Fig. 1: Phylogenetic relationships ofcertain munia finches. The birds inboldfacewere munias and mannikins (including sequenced from museum skins, with the date ofcollection indicated. © Biodiversity Heritage Library, http://www.biodiversitylibrary.org/; www.zoologicalbulletin.de; www.biologiezentrum.at Museum Collections as Sources ofGenetic Data 101 Cream-bellied mania Lonchiira pallidiventer was Payne 1999, 2001a,b, 2002), both museum skin spec- described as a species from a bird market in Jakarta imens and tissue samples were used as sources of (Restall 1996), the birds said to be from southern genetic material. By examining voucher specimens Borneo. The bird has not been seen in the field and is we confirmed or corrected the identification ofa num- thought to be a hybrid (van Balen 1998). Luis Bap- ber ofmolecular samples. TISTA provided feathers of two live birds that he These voucher specimens were prepared by the col- obtained from Robin Restall through the San Diego lector and retained by the museum as a document of Zoo. The bird was nearly identical in mtDNA sequen- identification. In most cases, the voucher specimen ce to chestnut munia Lonchiira (ferruginosa) atrica- was located, and examination showed it to be the pilla (1 bp different) and tricolored munia L. malacca species as identified by its genetic sequence, and not (3 bp different). The lack of difference from these as in the museum records. In one case the specimen munias suggests that L. ,,pallidiventer" is a hybrid, was not located and from the genetic results we sus- with the maternal parent L. (f.) atricapilla. The mito- pect the bird was misidentified (case 1): we were able chondrial gene is transmitted maternally, so the hybrid to idenfify the bird from the collector's measure- would have the mtDNA of its mother. Nuclear genes ments. The specimens were correctly identified for are needed to determine the fatherby molecular meth- most ofthe 126 tissue samples from six museums, but ods. VAN Balen (1998) suggested a hybrid origin with 8 ofthese birds (6 %) were misidentified. In cases (1, the parent species scaly-breasted munia L. piinctiilata 2 and 3) the cuckoo tissue was sequenced in another and white-bellied munia L. leiicogastra. Nevertheless, study (Johnson et al. 2000) and the incorrect identifi- Restall (1997) described the song oí pallidiventer" cation (in case (1), a new misidentification as „Cucu- as like the song of five-colored munia L. quinticolor lus vagans") was published and incorrectly entered Because munias learn their song from their father into GenBank. Our conclusions about avian systemat- (GüTTiNGER 1973; Clayton 1989), I. quinticolor^Na?, ics and evolufion would have been incorrect had we probably the father ofthe hybrid. not examined the specimens and field data. L. „pallidiventer" has a scaly feather pattern on the A Philippine cuckoo was identified as a hawk- 1 . flanks. The scaly pattern occurs in several species of cuckoo Cuculus (fugax) pectoralis and the record was munia though not in L.ferruginosa atricapilla and L. published with this identification. The genetic se- quinticolor. The pattern appears occasionally in L. quence from its tissue was unlike that ofother Cuculus malacca, which is an allospecies with L. ferruginosa fugax or C. pectoralis that we have sequenced, but it atricapilla (Restall 1997). Restall (1997) illus- was like oriental cuckoo C. saturatus. Neither the two trates several munia species that regularly have a North American museums that supported the field- scaly pattern of feathers or bars on the flanks, and work, nor the Philippine National Museum to which other species that sometimes have these patterns. The these museums directed us, had a register record ofthe scaly gene is found in many species that lack the pat- voucher specimen, said to be an unsexed spirit. The tern but is expressed in hybrids even those involving collector provided us with the wing measurement, 181 the silverbills L. castaneothorax, Euodice malabarica mm, indicating the bird was C. saturatus. aRnedstaE.llc(a1n9t9o7n)s o(bBsaeprtviesdtan,estibnuillitdti.,ng1inJuhlisy ca1p9t98i)v.e 2. A Philippine cuckoo identified as plaintive cuckoo Cacomantis merulinus yielded genetic sequence from „pallidiventer" but had no young, and Baptista had tissue the same as a Philippine brush cuckoo Caco- a pair of„pallidiventer" nest and lay but the eggs did not hatch. Hybrids are known for many estrildid mantis variolosus. Examination ofvoucher specimens species, including some in different genera and tribes showed that both were Cacomantis variolosus, one an and some intergeneric estrildid hybrids are even fer- adult female and one (the misidentified bird) ajuvenile. tile (Immelmann et al. 1977; Fehrer 1993). 3. Ajuvenile South African cuckoo identified as Klaas cuckoo Chrysococcyx klaas had a gene sequence like 3. THE IMPORTANCE OF MUSEUM SPECI- that of diederik cuckoo Chrysococcyx caprius and MENS AS VOUCHERS FOR IDENTIFICATION unlike another C. klaas. Examination of the voucher IN MOLECULAR GENETICS skin showed that the bird was ajuvenile C. caprius. In recent studies of the phylogenetic relationships in 4. A Bornean cuckoo was identified as oriental cuckoo cuckoos and in Old World finches (Sorenson & Cuculus saturatus lepidus. Sequence analysis of the © Biodiversity Heritage Library, http://www.biodiversitylibrary.org/; www.zoologicalbulletin.de; www.biologiezentrum.at & 102 Robert B. Payne Michael D. Sorensen tissue sample showed it to be like other Cacomantis hybridization. For market birds, captive birds in avi- variolosas. The bird was preserved as a study skin for culture, and wild birds caught and sampled for blood the collection of Sabah Parks. The Research Officer or feathers then released in the field, we recommend a (Zoology of Sabah Parks photocopied the specimen photograph for documentation. ) and its label with the data that matched the data ofthe Museum collections can provide archives of birds collector. The faxed photocopies showed the bird to used in genetic study when blood, tissues or feathers be ajuvenile Cacomantis variolosus. lost in the skinning process are preserved separately 5-9. Five South African ploceids were misidentified from the study skin, skeleton and spirit. Continued to species in a museum that has an active program of collecting is recommended (e.g., Remsen 1995; Winker 2000): new avian taxa are continually being collecting genetic samples and voucher specimens. discovered, our museums undersample the variation Examination ofthe voucher specimens gave identifi- ofbirds ofthe world, and we can sample the genetics cations that were consistent with the genetic sequence ofbirds of special systematic and conservation inter- information, so the birds had been misidentified by the collectors and the museum. est. Because not all specimens have been correctly identified in recent genetic studies, we recommend In another case a voucher specimen verified the that an active museum systematist be involved in museum identification where the identity was ques- genetics studies to identify the voucher specimens. tioned in the genetic results. Finally, we suggest that the importance of museum A number ofcountries require that specimens remain specimens as primary genetic resources be considered in balancing the views of molecular genetics use as in the country of origin as a condition of a permit to „destructive sampling" versus „added value" science. collect. This requirement can benefit the host country with specimens for scientific development and ACKNOWLEDGMENTS research, but it can make the specimens unavailable For providing access to their collections and permitting the for reference and repeated sampling in molecular sys- use ofgenetic samples, we thank AM (Auckland Museum, tematics (Ruedas et al. 2000). When specimens are Auckland, New Zealand); AMNH (American Museum of returned to the country oforigin, we recommend that Natural History, New York), BMNH (Natural History the museum take photographs and retain distinctive Museum, TriBnWg)Y; OBPBM (B.P. Bishop Museum, University ofHawaii); (National Museums ofZimbabwe, Bul- feathers with a museum registration number. For awayo); CAS (Chinese Academy of Sciences, Beijing); example, ZMUC saved the entire molted plumage of CMNH (Cincinnati Museum of Natural History, Cincin- a unique specimen of a Laniarius bush-shrike from nati); CU (Cornell University, Ithaca); DM (Durban Museum, Durban); DMNH (Delaware Museum ofNatural Somalia; when the bird was released to the field after History, Greenville); FLMNH (Florida State Museum of a long period in captivity (Smith et al. 1991), the Natural History, Gainesville); FMNH (Field Museum of entire set of feathers was retained as a permanent Natural Histoiy, Chicago); LSUMZ (Museum ofZoology, Louisiana State University, Baton Rouge); MCZ (Museum archival record at the museum. MNHN of Comparative Zoology, Harvard University); MVZ (Muséum National d'Histoire Naturelle, Paris); 4. DISCUSSION (Museum of Vertebrate Zoology, Berkeley); MSU (Michi- gan State University Museum of Zoology, East Lansing); A feather itself is both a sample for DNA sequence NMGL (National Museums and Galleries (Merseyside information and a voucher. Many species can be iden- Museums),NLMivMerpool); OU (Oklahoma University Stovall Museum); (National Museums ofMalawi, Blantyre); tified on the basis ofa single flight feather or distinc- QM (Queensland Museum, Brisbane); RMNH (Nationaal tive display feather, and museums can catalogue these Natuurhistorisch Museum, Leiden); UMMZ (University of with study skins. In our own studies, feathers from Michigan Museum of Zoology); UNAM (the Universidad Nacional Autónoma de Mexico); UNLV (University of avicultural sources were used as vouchers, such as the USNM Nevada at Las Vegas, Las Vegas); (National distinctive barred feathers of pictorella finch Hetero- Museum of Natural History, Washington, D.C.); UWBM munia pectoralis. Feathers removed from a museum (Burke Museum, University of Washington, Seatde); specimen can be returned to the museum and reat- WillFo,VZCal(iWfeosrtneira)n;FoYunPdMatio(nYaolfeVePreteabbroadtye ZoMoulsoeguy,m,CamNaerw- tached to the specimen as in the original skinning Haven); ZFMK (Museum Alexander Koenig, Bonn); and process, or labeled and stored in individual envelopes. ZMUC (Universitets Zoologiske Museum, Copenhagen). Blood samples from the same individuals that are UNAM, UWBM and Sabah Parks (Zoology) provided used for feather samples for mtDNA sequence might eirmsagfersomorlilvoeabnierddsspweecriemesennstnboywLuiinsthBeaiprticsotlale,cJtiüorngse.nFNeiactoh-- provide nuclear markers to resolve questions ofrecent lai and many aviculturists. Kristina M. Sefc helped deter- © Biodiversity Heritage Library, http://www.biodiversitylibrary.org/; www.zoologicalbulletin.de; www.biologiezentrum.at Museum Collections as Sources ofGenetic Data 103 mine molecular sequence data for Pholidornis and other Graves, G. R. & Braun, M. J. (1992): Museums: store- Old World songbirds. The National Science Foundation houses ofDNA? Science 255: 1335-1336. (NSF), Boston University and the University of Michigan Güttinger, H. R. (1973): Kopiervermögen von Rhythmus provided financial support. und Strophenaufbau in der Gesangsentwicklung einiger Lonchiira-Arten (Estrildidae). Z. Tieipsychol. 32: 374- LITERATURE 385. & & Hillis, D. M., Moritz, C. Mable, B. (eds.) (1996): Mol- Arctander, Dp.NAF.1ELDSÄ, J. (1994): Avian tissue collec- ecular Systematics, 2nd edn. Sinauer, Sunderland, tions for analysis. Ibis 136: 359-360. Massachusetts. 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(2001a): A single origin Massachusetts 02215, USA ofbrood parasitism in African finches: implications for ZOBODAT - www.zobodat.at Zoologisch-Botanische Datenbank/Zoological-Botanical Database Digitale Literatur/Digital Literature Zeitschrift/Journal: Bonn zoological Bulletin - früher Bonner Zoologische Beiträge. Jahr/Year: 2003 Band/Volume: 51 Autor(en)/Author(s): Payne Robert B., Sorenson Michael D. Artikel/Article: Museum Collections as Sources of Genetic Data 97-104