Juan Cristóbal Gundlach’s collections of Puerto Rican birds with special regard to types

The German naturalist Juan Cristóbal Gundlach (1810–1896) conducted, while a resident of Cuba, two expeditions to Puerto Rico in 1873 and 1875–6, where he explored the southwestern, western, and northeastern regions of this island. Gundlach made repre­ sentative collections of the island’s fauna, which formed the nucleus of the first natural history museums in Puerto Rico. When the natural history museums closed, only a few specimens were passed to other institutions, including foreign museums. None of Gund­ lach’s and few of his contemporaries’ specimens have survived in Puerto Rico. We lo­ cated 191 bird specimens (43 species) collected there by Gundlach, all of which are in foreign institutions, especially Museum für Naturkunde Berlin. Here we list all located specimens and include data associated with them. Six new species were described out of the Gundlach collections from Puerto Rico, three of which are still taxonomically recog­ nized today. Information about the types of those taxa is given.


Introduction
Dr. Juan Cristóbal (Johann Christoph) Gundlach (1810Gundlach ( -1896 arrived in Cuba from his native Germany in Janu ary 1839, intending to stay only a short time. Instead, he from the western part he explored the northern region to San Juan, as well as the south western region of Puerto Rico. Additionally he made two trips to the central parts of the island (Lares and Gaguana/Jayuya).
His zoological interests were manifold, and he pub lished his observations widely, not only within Cuba, but also in several international journals, in Spanish, Ger man, and English. Gundlach amassed superb represen tative collections especially insects, molluscs, reptiles, mammals, and birds. He maintained an active exchange of specimens with foreign institutions and friends, so a considerable part of his collection was dispersed among different countries from the very beginning. This result ed in single specimens collected by Gundlach housed in several diverse collections today. Our purpose here is to list those vouchers of the avifauna of Puerto Rico avail able for research on this island as well as for international taxonomists. We also hope to stimulate interest of muse um curators to search for additional specimens in their collections.
A summary of Gundlach's lifework, especially of his two expeditions (including a map) as well as his influence on the development of natural history in Puerto Rico is presented in Wiley et al. (2014).

Collections of Puerto Rican Birds made by Gundlach and his colleagues
Even as a boy, Gundlach was interested in studying ani mals and got his introduction to taxidermy by helping his older brother preserve anatomical and biological speci mens. After his university studies, Gundlach decided to make an expedition to Suriname. To finance the travel, Robert Wilhelm Bunsen (1811-1899), second director of the "Verein für Naturwissenschaften Kassel", initiated an arrangement wherein shares were issued to support Gund lach's 1838 expedition to Suriname. Some 105 sharehold ers subscribed for 203 shares with a value of 1218 Thaler (Dathe and Gonzales Lopez 2002). This financial support of Gundlach's expedition was made on the condition that he sends scientific objects back to Germany to reimburse the travel expenses. Although Gundlach made it only as far as Cuba, he sent large shipments of scientific materi al to Germany (Anonymous 2011; Dathe and Gonzales Lopez 2002). Eduard Sezekorn (1796-1869), managing director of the Verein für Naturwissenschaften Kassel, or ganized the sale of the collected materials.
This was the start of Gundlach's intensive longterm collecting activities in the Caribbean. The specimens gathered there were widely distributed from the very be ginning. But Gundlach also retained extensive collect ed materials for his own museums of natural history in Cuba. In his autobiography, Gundlach (1896) wrote that one specimen of Mellisuga helenae which he collected in 1844 was the first specimen of his personal collection. He realized that the hummingbird was an unknown species and therefore he decided to keep it rather than to send it to Germany. Thereafter, Gundlach kept one specimen of each species, except of fish and large reptiles, for his own collection (Gundlach 1896). A third part of his collected specimens was sent to his friends and colleagues in Cuba as well as all over the world for exchange or to serve as the basis for scientific discussions (e.g. Ramón M. Forns (La Habana, fl. 1858), Tomas Blanco (San Juan), Agustin Stahl (Mayagüez), George N. Lawrence (New York), and Wilhelm Peters and Jean-Luis Cabanis (Berlin)). In prin ciple, the specimens which Gundlach collected in Puerto Rico were intended for comparison with the specimens from his Cuba collecting and suffered the same fate as the specimens from Cuba. By the 1870s, Gundlach's travel ex penses should have been recouped by his investors, so the Puerto Rican specimens remained in Gundlach's collec tion or were sent only to friends and scientific colleagues, and none should have been sent to Germany to settle his 1838 account. Finally, in gratitude for their patronage and benefaction over the years, Gundlach gave many of his bird specimens to friends as salon decorations.
Today, Gundlach's personal collection is housed in the Instituto de Ecología y Sistemática (IES, La Habana, Cuba). The IES collections have received little evaluation since the catalogues of Gundlach (1895) andValdes Ragués (1914), with the exception of recent reviews of Cuban birds (Aguilera Román and Garrido 2000;Aguilera Román et al. 2002;Wiley et al. 2008), reptiles and amphibians (Moreno García et al. 2002), and Noctuoidea (Lepidoptera) (Beck er 2002). We examined 18 Cuban collections (Wiley et al. 2008), but found bird specimens from Gundlach's Puerto Rican expeditions only at the IES, although several of his Cuban bird specimens were dispersed among other Cuban institutions. Most of the specimens collected in Puerto Rico by Gundlach were sent to institutions in the United States and Europe, especially Germany. We found Puerto Rican specimens collected by him in seven institutions (Table 1; see Suppl. material 1 for names of all institutions we visit ed or queried regarding holdings of Gundlach's Puerto Ri can bird specimens). Most of these specimens were direct donations from Gundlach, whereas a few arrived at those institutions as part of the collections of others, such as the Henry W. Bryant collection containing specimens given by Gundlach to G. N. Lawrence. Most of the Gundlach specimens received by the U. S. National Museum of Natural History (USNM) came as direct donations (~12 separate acquisitions) from him. There was one accession in 1868 of ten specimens from Puerto Rico, and an accession in 1877 of 9-10 birds from the West Indies (in litt. James Dean to JWW; 10 Novem ber 2006; USNM).
Today, the most comprehensive collection of Gund lach's specimens from Puerto Rico is housed in the Mu seum für Naturkunde Berlin. Once he settled in Cuba, Gundlach sent his ornithological observations to Eduard Sezekorn, who transformed Gundlach's notes into manu scripts and sent them to Jean Louis Cabanis (1816Cabanis ( -1906, the bird curator of the Zoological Museum Berlin (ZMB, today Museum für Naturkunde). Cabanis published the articles with remarks in his Journal für Ornithologie. The first specimens from Cuba arrived at ZMB in 1862 through Sezekorn and were most probably part of the re muneration of Gundlach's travel expenses to the "Vere in für Naturwissenschaften Kassel". A direct relation between the ZMB (Wilhelm Peters, Jean-Luis Cabanis) and Gundlach was established in 1861 (Peters) and 1862 (Cabanis) at the latest, more than 10 years before his first trip to Puerto Rico (Museum für Naturkunde, Historische Bild u. Schriftgutsammlungen, Zool. Mus. [hereafter MfNHBSZM], Signatur ZMB S I, Gundlach, I.). Follow ing his participation in the Exposition Universelle in Par is, Gundlach visited the ZMB in 1867 for a week, where he met with the museum's curator Wilhelm Peters (1815Peters ( -1883. Further specimen donations to the ZMB followed (MfNHBSZM Signatur ZMB S I, Gundlach, I) The material collected by Gundlach in Puerto Rico was sent in three shipments to the ZMB (arrival dates March 1874, June 1874, October 1876). Birds from other collec tors (Blanco, Krug) in Puerto Rico were included in those shipments to the ZMB (in litt. JCG to Wilhelm Peters [hereafter WP]; 15 February 1874; from Fermina, Cuba; MfNHBSZM, Signatur ZMB S I, Gundlach, I., p. 219a). All, however, were registered with Gundlach as collector, and the determination of the actual collector is difficult or impossible today because the original labels were not conserved. Further, it seems that not all specimens Gund lach sent were maintained in the Berlin collection. Thus, a hummingbird mentioned by Gundlach in a letter to Pe ters dated 15 February 1874 is not available today and could not be traced in museum catalogues (MfNHBSZM, Signatur ZMB S I, Gundlach,I.,p. 129a). This, and other specimens, may have been lost, destroyed, or exchanged with other collections.
The first of Gundlach's three shipments included 34 bird specimens that arrived at ZMB in March 1874. A second shipment, with 28 birds (according to the cata logue) arrived in June 1874. All of those birds were col lected during Gundlach's 1873 expedition to Puerto Rico. The month of collection was recorded for most birds in the second shipment, whereas it was not for most speci mens in the first shipment.
A third shipment of three boxes (including one of birds) of specimens collected during Gundlach's 1875-76 expe dition to Puerto Rico arrived at ZMB in October 1876. Gundlach mentioned that among these birds were speci mens from Krug's collection (in litt. JCG to WP; 17 Au gust 1876; from Mayagüez, Puerto Rico; MfNHBSZM, Signatur ZMB S I, Gundlach, I., p. 259.). Cabanis con firmed that 30 stuffed birds, 67 eggs, and 7 nests arrived in October 1876 (MfNHBSZM, Signatur ZMB S I, Gund lach, I., p. 266). Twentyeight of those birds were cata logued in October 1876. After this date seven additional specimens from Puerto Rico arrived in June 1878, January 1879, December 1880, and June 1881. Those specimens were probably part of Stahl's collection, which Gundlach had received at that time (in litt. JCG to Jean Louis Cabanis; 29 September 1878; MfNHBSZM, Signatur ZMB S I, Gundlach,I.,p. 294). It cannot be excluded, however, that Gundlach sent some specimens of his own expeditions which had been maintained in his personal collection.
Upon their arrival in Berlin, the specimens were as signed entrance numbers (Bnumbers) and, after prepa ration, inventory numbers. Only five specimens of the second shipment (1874) were inventoried later. All of Gundlach's specimens for the ZMB collection were mounted when received. Collectors labels were removed usually and new museum labels were created. Of the inventoried specimens, 70% have survived (98% of the eggs), and 85% of those have been transformed to skins. The transformation from mounts to skins was initiated by Erwin Stresemann  in the middle of the 20 th century to better protect the specimens.
In general, Gundlach seemed to collect for taxonomi cal purposes only, and he used the specimens as vouchers and for determination. But the exact collecting locality and the collecting date are never mentioned. For some specimens a month is given on the label as Gundlach mentioned that several birds are to be found in different seasons only. The lack of more data is unfortunate, be cause locality and date data could provide a more useful baseline in assaying what has been the dramatic change of biodiversity that has occurred in Puerto Rico, including several species that have been extirpated from in the is land or are now extinct (Wiley 1985, Snyder et al. 1987).

Puerto Rican Bird Specimens Collected by Juan Gundlach
We present an annotated inventory of Gundlach's bird specimens (skins and mounts, as well as eggs and nests) from Puerto Rico in the two largest collections: the col lection at the MfN, Berlin, and the collection at the IES, La Habana, with notes on Gundlach's Puerto Rican spec imens encountered in other institutions. Both the IES and MfN specimens have been unreported, except for Valdes Ragués' (1914) incomplete catalogue of the Museo cuba no "Gundlach" when it was housed at the Instituto de Se gunda Enseñanza, La Habana. The IES specimens were "discovered" among Gundlach's collection of Cuban skins, mounts, eggs, and nests during our recent invento ry. The MfN specimens remained more or less unknown or international scientists had thought they had been de stroyed in WWII. Our 2011 inventory, however, revealed that although the MfN collections were damaged during the Allied bombing of Berlin, many of Gundlach's spec imens and their records survived. Figures 1 to 3 are ex amples of how the specimens appear today in the MfN.
The list, as well as the species names (Latin, English) including current subspecies names, follow Dickinson and Remsen (2013) and Dickinson and Christidis (2014). The scientific name used by Gundlach (1874Gundlach ( , 1878a is add ed in brackets. For each species, institution and catalogue number(s), sex, date(s) of collection, and locality, as avail able, are presented. As already written, none of the IES Puerto Rican specimens has detailed data. Nevertheless, some of the specimens can be matched to Gundlach's pub lished accounts of his collecting activities, and thus fur ther data were derived from interpretation. For specimens housed in MfN, we use the original institution name of ZMB. The ZMB specimens could generally be assigned to the different collecting expeditions due to the accession date. We present information included by Valdes Ragués in his 1914 catalogue of Gundlach's collection in the Insti tuto de Segunda Enseñanza. Puerto Rican specimens in the Instituto de Segunda Enseñanza museum were identified by a yellow label, but Valdes Ragués (1914) did not always mention that information in his catalogue. Full names and acronyms for institutions are presented in Table 1. Ex panded specimen data are available from the authors.
Following the specimen list, we provide a list of the types belonging to the Gundlach collection from Puerto Rico. Six species were described from that collection. Ex cept for Asio portoricensis, all of those species were de tected by Gundlach himself, but in some cases they were "officially" (in the sense of the ICZN) published by other authors. This was true for G. N. Lawrence (New York), who received a bird collection from Gundlach and J. L. Cabanis (Berlin).

List of specimens
The following list summarizes the information on the specimens of Puerto Rico originally belonged to the collection of Gundlach (Gundlach mainly given as collector).
The catalogue is structured as follows:

Margarops fuscatus fuscatus, Pearly-eyed Thrasher [Margarops fuscatus]
It is remarkable that this nowabundant and obvious spe cies (Snyder et al. 1987;Arendt 2006) was rare in Puerto Rico during Gundlach's time. He wrote (1878a): "Sola mente en dos ocasiones he observado esta especie, y la creo poco común, porque pocas personas la conocían." He collected one in a coffee plantation, and observed a pair near Utuado in July. Type series. Lawrence (1875) described this species and attributed its discovery to Gundlach, but Gundlach (1878b) corrected this, stating that the specimen was col lected by Tomás Blanco y González in Puerto Rico. Gund lach most probably received the specimen from Blanco in 1868. He later transferred it to the Smithsonian Institution in Washington, D. C. (today USNM) and there Gund lach was given as collector. As Lawrence (1875) wrote "Type in National Museum, Washington", the type series con sists of only one specimen which is the only available specimen of that species at the USNM "collected by Gundlach" and is therefore the holotype. An additional specimen of this species from the Gundlach collection is housed at the ZMB (ZMB 24887), but it was collected by Stahl between 1878 and 1880 and was sent directly from Gundlach to Berlin. It is therefore not part of the type series. LeCroy (2012) noted that a further specimen at the American Museum of Natural History (AMNH 3262) has a type label for Chlorospingus speculiferus. But she did not consider it as a type specimen because it is not dated and Gundlach is not indicated clearly as the collector. We follow this argument. It appears that Gundlach never col lected this species himself. Remarks. The specimen was catalogued at the USNM on 18 October 1878, but Lawrence previously wrote in the description in 1875 that the specimen was housed in the National Museum, Washington. We do not know when the specimen arrived at the USNM. It may be that it remained there uncatalogued until the discussion about the type with Gundlach started in 1877. As it is the only specimen of this species collected by "Gundlach" at the USNM, we believe that USNM 75331 should be the type specimen.

Now. Contopus latirostris blancoi
Type series. Not specified by Cabanis (1875). There is only one specimen noted in the inventory catalogue of the ZMB (ZMB 21492). But because the species was dis covered and named by Gundlach all the specimens col lected during the first expedition to Puerto Rico should be regarded as types according the ICZN (1999). Anoth er specimen collected by Gundlach is housed at the IES (IES 2576). The date is given as "December" on the label. As Gundlach left Puerto Rico on 4 December 1873 after his first expedition to Puerto Rico it is more likely that he collected this specimen during his second expedition Remarks. Gundlach discovered this new species and named it in honor of his friend Tomás Blanco y González (1840-1892, living in Puerto Rico. Gundlach (1874) in cluded the species in his published list without descrip tion. Cabanis (1875) added the missing description, so he is officially the author of this species, following the ICZN (1999).

Now. Megascops nudipes nudipes
Type series. Not specified by Gundlach (1874). There are two specimens noted in the inventory catalogue of the ZMB. Therefore these are syntypes.  [missing, fide Valdes Ragués (1914)]) and the AMNH (AMNH 44792-44795) which were (most probable for the IES and two of the AMNH specimens) collected during Gundlach's second expedition to Puerto Rico or even later by Agustin Stahl . Thus, due to missing exact information we do not regard any of these specimens as types.

Now. Asio flammeus portoricensis
Type series. Ridgway (1882) attributed the discovery of this species to Baird et al. (1874) when the existence of a new species of this genus from Puerto Rico was de termined. Lacking further material for comparison, this species remained unnamed in 1874. In his description, Ridgway (1882) wrote that he had studied 4 specimens from Puerto Rico, one specimen in the USNM (USNM 39643 which was illustrated and described as aberrant by Baird et al. (1874) and three specimens which were col lected by Gundlach and received for determination from George Newbold Lawrence. Ridgway reported that all four specimens were very similar and described the new species. Therefore, all four specimens are syntypes. Some of Gundlach's specimens passed to Lawrence

Now. Chlorostilbon maugaeus
Type series. Not specified by Gundlach (1874). In 1878, Gundlach (1878b) wrote that he collected one male of this new species which was sent to Berlin later and so it is the holotype.

Remarks.
Gundlach was not aware of the description of Sporadinus maugaeus Viell. 1817 when he described Chlorestes gertrudis. In his second publication on the birds of Puerto Rico (1878b), Gundlach suspected synon ymy with this species but he was uncertain.
Five additional specimens of this species were collect ed by Gundlach (ZMB 22672-22674;IES 2578;AMNH 38784), but all have been collected during his second ex pedition to Puerto Rico and, therefore, we do not regard them as types.

Now. Psittacara chloropterus maugei
Type series. Cabanis mentioned in his description as well as in Cabanis (1881b) that he described the species based on two left wings, which means his description is based on two specimens. Remarks. Three wings of a parakeet from Isla de Mona were collected by Dr. Claudio Federico Block (or Bloch), a Danish physician and hunter from Mayagüez, who gave the wings to Gundlach probably in 1875. Two of the wings arrived in Berlin in October 1876 (B 14144). But they did not get inventory numbers and the whereabouts of the wings are unclear, so they are apparently lost. The wherea bouts of the third wing is unknown, too (Olson 2015).

Summary
In summary, we can account for 191 bird skin and mount specimens collected by Gundlach in Puerto Rico, repre senting 43 species. An additional 69 eggs (19 species) and 6 nests (5 species) were located, including eggs of 7 species not represented by skins. A total of 147 skin and mount specimens collected by Gundlach in Puerto Rico were found in foreign institutions, with the Muse um fuer Naturkunde holding the largest number (102), including two holotypes and two syntypes ( Table 1). The MfN collection contains 53.4% of all specimens, and 93.0% of the species in all collections examined. Further, MfN specimens represent 26.1% of the 153 species Gundlach reported from Puerto Rico (Gundlach 1878a(Gundlach , 1878b. We found 44 specimens (41 extant), representing 28 species, collected by Gundlach in Puerto Rico in the IES collection. Valdes Ragués (1914) listed 18 Puerto Rican specimens by name or group in his analysis of Gund lach's collection, with another improbable 15 Audubon's Shearwaters. The IES collection contains 23.0% of all specimens, and 65.1% of the species in all examined col lections (Gundlach 1878a, 1878b; Table 1). Further, the IES specimens represent 18.3% of the 153 species Gund lach reported from Puerto Rico.
Conspicuously missing from all collections of Gun dlach's bird skins from Puerto Rico are specimens of shorebirds, waterfowl, and waders, even though he ac tively collected in several coastal and wetland areas and did take several eggs of waterbirds.
Unfortunately, none of Juan Gundlach's journals or field notes has been found. Such material might provide muchdesired additional data on Gundlach's collected specimens, much of which would be valuable in further determination of types. Some information may be avail able in Gundlach's correspondence, and we urge others to search for all such materials.

Concluding remarks
The theft of the Puerto Rican Parrot specimen, sadly, was not the last of Gundlach's bird specimens to have been stolen from the IES collection. In 2007, thieves again raided the collection, this time making off with sever al Gundlachcollected birds. Tragically, the only Cuban Macaw (Ara tricolor) specimen in Cuba (and one of 19 known specimens worldwide; Wiley and Kirwan 2013) was taken by the thieves, who also stole air condition ers and other valuable items essential for maintaining the collections. The specimens were most likely taken not for their scientific value or for their value to rogue pri vate collectors, but rather for use in local spiritual rituals. Although substantial improvements have been made in security, the IES collections are still vulnerable to future raids. The IES collection of birds and other natural histo ry specimens is an important record of Cuban and Puerto Rican biodiversity, but is threatened by a lack of funding to maintain the collections. Such funding is desperately needed to prevent the further degradation or loss of these important treasures.
We hope to stimulate interest of museum curators to search for additional specimens in their collections.