Catalogue of the Vascular Plants of Ecuador

By Peter M. Jørgensen

The following overview of the history of collecting in Ecuador, while not exhaustive, does highlight the most important collectors and collecting expeditions. It also provides some data that are not readily accessible. The sources for this account are Diels (1937), Acosta-Solís (1969b), Holm-Nielsen (1986a), Molau (1986), Wiggins and Porter (1971), Jørgensen et al. (1992), and Renner (1993), as well as several biographies and general references (Stafleu & Cowan, 1976–1988; Stafleu & Mennega, 1992–1997; and Barnhart, 1965). The author's observations from personal knowledge are also included. Herbarium acronyms follow Holmgren et al. (1990).

The 18^th century — The first botanist to collect plants in Ecuador for scientific purposes was Joseph de Jussieu (1704–1779), from France, a participant in the French Geodesic expedition to Ecuador from 1735 to 1743 (Zúñiga, 1977). The primary objective of the expedition was to measure the length of a degree close to the equator. The length of a degree would be constant if the earth was a perfect sphere, so a similar expedition was simultaneously making measurements in Lapland, Sweden, for comparison. This seems a trivial question today, but it was very important to cartography and navigation in the 18^th century. Jussieu's first collections were made in Panama, and he continued collecting in Ecuador for the duration of the eight-year expedition. After the successful completion of the expedition, he continued to collect in Peru and Bolivia and remained in South America until 1771. However, Jussieu went insane while working in Ecuador (von Hagen, 1945). A shortage of money kept him from returning to Paris at the end of the expedition; later, his medical abilities may have kept him in Quito where his services were needed during a smallpox outbreak (Steele, 1964). Linné (see Steele, 1964) wrote, "Those who have been with him say that he has done almost nothing [of a botanical nature], he has only practiced [medicine]." A contributing factor to Jussieu's insanity may be found in the fact that he lost most of the documentation for his work; he had entrusted a trunk full of notebooks and dried plant specimens to a servant who ran away with the "treasure" across the border to Brazil. A minor portion of his collections survived and were brought to France (P), where Lamarck used them in the elaboration of his famous Encyclopédie Méthodique (1783–1823).

Pedro Franco Dávila (1711–1786), born in Guayaquil, was a collector of natural history specimens. He lived in Paris from 1740 to 1771 where he bought the specimens that constituted his personal cabinet (Calatayud, 1988). After several letters to King Carlos III of Spain, in which Dávila offered his cabinet to the king, his offer was finally accepted in October 1771. Dávila established, with his collections, the Real Gabinete de Historia Natural, of which he was the first director (Calatayud, 1988). The museum was not opened to the public until November 1776. Dávila wrote a guide to collecting plants and other items from nature. This guide, which was distributed to all officials of the crown in Latin America, was one of Dávila's achievements in the documentation of the natural richness of the Spanish possessions in Latin America.

Thaddäus Peregrinus Xaverius Haenke (1761–1816), a Czech (Austria–Hungary), and Luis Née (1734–1807), a Frenchman, were both botanists on Alejandro Malaspina's expedition around the world from 1789 to 1794. Antonio de Pineda, a Guatemalan zoologist, was in charge of the scientific part of the expedition (Cerezo, 1994). Haenke was a pupil of Josef Gottfried Mikan in Prague and later of Nicolaus Joseph von Jacquin in Vienna. He was widely respected in central Europe because he edited the eighth edition of Linné's Species Plantarum (1791), a work he must have almost finished before leaving on the Malaspina expedition. He was invited to participate in the expedition by King Carlos IV. Haenke arrived a few hours late at Cádiz, the port of departure of the expedition. To catch up, he sailed with a merchant to Río de La Plata where his ship sank; he only managed to save his royal letter of appointment and a few other personal items. From La Plata, he traveled overland to Santiago, Chile, where he finally met with Malaspina (Ibáñez, 1993). The expedition arrived at Guayaquil on 1 October 1790. During the 28 days the expedition was in Guayaquil, Haenke collected only in the surroundings of Guayaquil, Daule, and Taura (Muñoz, 1993). On the way back to Spain he was ordered, but probably not against his will, to travel overland and collect from El Callao in Peru to Buenos Aires. He never reached Buenos Aires, but settled in Cochabamba, Bolivia, where he died (Ibáñez, 1993). Haenke had, during the entire expedition, sent dried plants to a private company, Hieke, Rautenstrauch, Zincke und Ko., and it is assumed that these collections arrived in Prague, where Carl Borivoj Presl and Jan Svatopluk Presl worked with the collections at PR and PRC, respectively. The other set (or sets) of duplicates that arrived in Cádiz with Malaspina were never incorporated into any extant herbaria, and the only trace of these plants is a report from the "Depósito Hidrográfico" listing 85 bundles of plants (Muñoz, 1993). Haenke collected approximately 15,000 numbers during the expedition. C. Presl published the new species in Reliquiae Hankeanae (1825–1835). Upon his death, his notebooks and other materials were sent from Cochabamba to Madrid where they remain at MA; for other deposits of documents, see Ibáñez (1993).

Prior to the expedition Née had collected widely in Spain, and that is probably why Pineda elected him to participate in the Malaspina expedition. Née made a trip to Chimborazo and Tungurahua; localities visited were Las Bodegas de Babahoyo, El Caracol, Pozuelo, San Antonio, Guaranda, Chimborazo, El Arenal, Mocha, Pelileo, Baños, Tungurahua, Río Filca, and Río Caluma (Muñoz, 1993). This itinerary covers a vertical climb of maybe 5,000 m, and Née covered almost 500 km in 18 days (28 km/day). He later promised, in a letter to Mutis, that he would never again make such hasty trips. During this "excursion," Née made about 200 collections; his total was about 12,000 collections for the entire expedition. Most, if not all, of his collections from this expedition are deposited in Madrid (MA). Née was, like Haenke, ordered to cross South America collecting on foot. He went from Concepción north to Santiago and then east to Buenos Aires where he again caught up with Malaspina. For further information, see Muñoz (1993). Haenke and Née did not seem to collaborate during the expedition. A certain level of competition must have existed among them, which may explain Née's lack of concern for the Haenke collections that arrived in Spain.

Juan José Tafalla (1755–1811), from Spain, worked with Ruiz and Pavón in Peru from 1785 to 1788, when Ruiz and Pavón left for Spain. Tafalla continued working in Peru and sent material back to Madrid. He took part in the founding of the San Marcos University in Lima and also founded the botanical garden in Lima. In 1799, he was ordered to undertake an expedition to Guayaquil (Estrella, 1991). The expedition was composed of Tafalla, chief of the expedition, Juan Agustín Manzanilla (a botanist who arrived from Spain in 1793), José Gabriel Rivera (artist), and Xavier Cortés (artist). The expedition spent the first three years, 1799–1802, in the coastal areas around Guayaquil evaluating the quality and quantity of numerous wood products used for ship building (Estrella, 1991). Between 1803 and 1804, they collected in the Andes of Ecuador, particularly around Cuenca and Loja. This part of the expedition was spent mainly on quinine research (Estrella, 1991). Many of Tafalla's observations and collections were the basis for several of Ruiz and Pavón's publications on this subject between 1792 and 1826 (Estrella, 1991). Tafalla's collections are often attributed to Pavón and are deposited in Madrid (MA); because of insufficient labeling it is often difficult to ascertain if they were actually collected in Ecuador and by Tafalla and Manzanilla. Tafalla died in Lima in 1811, and his work Flora Huayaquilensis remained unpublished until Estrella (1991) found a document that explained how to separate Tafalla's plates from those of Ruiz and Pavón.

The 19^th century — Very little is known about the first genuine Ecuadorian botanist, José Mejía del Valle y Lequerica (1775–1813), and no known collections remain. It is known that he sent some material to José Celestino Bruno Mutis in Bogotá and that some of his collections may remain in the Mutis collections in Madrid (MA). Most of his collections were made near Quito, but he also collected in the coastal lowlands at Santo Domingo and Atacames. A single handwritten manuscript has been found, titled Plantas Quiteñas, along with several letters (Estrella, 1988). Mejía left Ecuador in 1806 and took up politics in Spain, where he died in Cádiz from yellow fever.

Even less is known about his teacher in botany, Anastasio Guzmán (?–1807), from Spain, who arrived in Quito in 1801 via Buenos Aires, Chile, and Peru (where he spent some time with Tafalla). Guzmán was a pharmacist from Seville who came to Ecuador to study chemistry and nature, and he financed the trip and his research with personal funds (Estrella, 1988). He died during an expedition to Llanganates in search of the "lost" Inca gold. Mejía, and later his widow, tried to gain possession of Guzmán's materials, but they were unsuccessful. According to Estrella (1988), some of his materials are still conserved in Quito. Guzmania Ruiz & Pav. and Ranunculus guzmanii Kunth were named after him. Tafalla must have suggested the generic name to Ruiz and Pavón, as it is unlikely that they ever met Guzmán.

In 1802–1803, Friedrich Wilhelm Heinrich Alexander von Humboldt (1769–1859), from Germany, and Aimé Jacques Alexandre Bonpland (1773–1858), from France, were the first botanists to make a significant collection of Ecuadorian plants that was successfully brought to Europe. Their collections were deposited in the major herbaria (P and B) and led to the publication of numerous new Ecuadorian taxa. Karl (Carl) Sigismund Kunth worked on the Humboldt and Bonpland collections, and he described most of the encountered new species in Humboldt, Bonpland, and Kunth's Nova Genera et Species Plantarum (1815–1825). Humboldt obtained permission from the Spanish king to travel in Latin America and left Europe from Cádiz. In brief, Humboldt and Bonpland arrived at Bogotá in July 1801, where Mutis received them. At his suggestion they took Francisco José de Caldas along with them to Quito (von Hagen, 1945). They left Bogotá in September and arrived in Quito on 6 January (Sandwith, 1926; von Hagen, 1945). In Quito, Humboldt and Bonpland met the aristocracy and, among them, the young Carlos Montúfar, who accompanied them throughout the rest of their journey (Botting, 1973). They must have also met with Guzmán. They spent a total of six months in Quito and surrounding areas. A complete itinerary for Ecuador and Peru can be found in Sandwith (1926). They left Ecuador from Loja, and on their way to Mexico from Lima they stopped in Guayaquil where they collected with Tafalla in January and February, 1802, before finally leaving Ecuador (Sandwith, 1926; Botting, 1973; Estrella, 1991). Humboldt had, in part through the works of Kunth and, to a certain extent, Bonpland, a significant impact on systematic botany. Equally important was his influence on plant geography; his illustration of the plants growing on the slopes of Chimborazo, reproduced on the cover, played a significant role in that context.

Francisco José de Caldas (1771–1816), from Colombia, arrived in Ecuador in January 1802 and remained there until 1805. It is unclear how many plants Caldas collected, but he described Ullucus tuberosus in 1809, apparently based on material he collected in Ecuador. He traveled extensively in Ecuador and went as far south as Loja. Whether Caldas did not get along with Humboldt, or Humboldt was afraid of the intellectual competition from Caldas, as suggested by Murillo (1951, 1960), is unclear. Caldas, who had invented the hypsometer—a way of measuring the elevation by measuring the temperature of boiling water—was never part of the Humboldt and Bonpland team as was Montúfar. The number of collections made by Caldas in Ecuador is unknown. His collections may have been incorporated into Mutis's collections; equally probable is their complete destruction around the time when Caldas took part in the revolutionary movement and was arrested and executed in Bogotá.

The second very important set of Ecuadorian collections was made by William Jameson (1796–1873), an Englishman, who lived in Ecuador with minor interruptions from 1820 until his death. He spent the first six years in the area of Guayaquil, but after 1826 most of his collecting took place in the highlands of the Andes. Jameson taught botany and chemistry at the university in Quito. His collections are preserved in London (K and BM), but many duplicates have been distributed to a large number of institutions. Jameson's most important publication was Synopsis Plantarum Æquatoriensium (1865). Unfortunately, many of his collections do not include a precise location. A reason for this was given by Spruce, who wrote, "[Jameson] has had a drunken (and worse) wife hanging on him for forty years, who burns his dried plants, whenever she can get hold of them, so that he can keep no herbarium." However, many species have information on locality in Synopsis Plantarum Æquatoriensium, so a careful matching of the vouchers with his book may reveal more information than first meets the eye. Acosta-Solís (1969b) persuaded the Central University to reprint Jameson's Synopsis in 1940, using the argument that the second volume was left unpublished. The copy at MO is complete, however, and the second volume was a presentation copy from A. Deiting of Guayaquil to R. Spruce; the dedication is dated 25 July 1887.

Francis Hall (?–1834), from England, collected with Jameson and Bousingault (a French geologist) in 1831–1832, mainly in the Andean provinces of Pichincha, Cotopaxi, Napo, Tungurahua, and Chimborazo. His observations were published in a paper titled Excursions in the neighbourhood of Quito and towards the summit of Chimborazo (Hall, 1834). Hall's descriptions of the surroundings of Quito are precise, and it is obvious that few changes have occurred in the native plant composition of the fences and hedges in the inter-Andean area in the past 170 years. The continuation of the paper (Hall, 1835) indicates the year in the title to be 1830, but the mistake is corrected in the second part. Hall's collections were deposited at K. He died in Quito.

Richard Brinsley Hinds (1812–1847), George W. Barclay (dates unknown), and Andrew Sinclair (?–1861), all from England, took part in the voyage of H.M.S. Sulphur and Harrier around the world. The collections from that trip are housed in London (BM). It was not among the main goals of the journey to make botanical collections. The Sulphur's orders (Belcher, 1843), dictated, "Large collections of natural history cannot be expected, but … the medical officers will no doubt be anxious to contribute their share to the scientific character of the survey." Belcher (1843), captain for most of the voyage, described the journey, but placed very little emphasis on the scientific aspect and hardly even mentioned the scientists on the expedition. The last third of the second volume of Belcher (1843) is an article by Hinds (1843) in which he provides an overview of The regions of vegetation. Staffleu's (1968) note on the dates of publication in the facsimile version of Bentham's The Botany of the Voyage of H.M.S. Sulphur (1844–1846) mentions only Hinds and Barclay as collectors of the botanical specimens. He failed to note, however, that Bentham (p. 182) indicated that Sinclair also collected material on which some of the many new species were based. Bentham listed the following localities in Ecuador: Puná, Guayaquil, Batahoya [Babahoyo], Bodegas, Salango in Colombia (which undoubtedly refers to the Island of Salango on the Ecuadorian coast), Punta Santa Elena in Colombia (also undoubtedly in Ecuador), and Atacames. The visit to Ecuador lasted only slightly over a month from about 6 September until about 12 October 1838.

The Galápagos Islands were discovered by Tomás de Berlanga in 1535 (Slevin, 1959). Plant collecting started almost 300 years later and progressed very slowly from 1825 to 1846. These collections were all included in Joseph Hooker's work from 1847 dealing principally with Darwin's collections (Hooker, 1847; Porter, 1980). David Douglas (1798–1834) and John Scouler (1804–1871), both from England, made the first 40 collections on 10 January 1825; only 18 survived to be treated by Hooker. James Macrae (dates unknown), from England, made 41 collections on the island of Isabela between 26 March and 2 April 1825; 37 were included by Hooker (1847) and 20 represented new species. Hugh Cumming (1791–1865), from England, made nine collections in 1829. Charles Darwin (1809–1882), also from England, was on the Galápagos from 15 September to 20 October 1835. He collected 209 plants on Española, Isabela, San Cristóbal, and Santa María Islands; Hooker (1847) described 78 as new. Porter (1980) provides information on where the type material is located. Abel Aubert du Petit-Thouars (1793–1864) and Adolphe-Simon Neboux (dates unknown), both from France, visited the Galápagos from 21 June to 15 July 1838; they collected only eight numbers.

Thomas Edmonston (1825–1846) and John Goodridge (dates unknown), both from England, collected 41 numbers on Galápagos between 6 and 16 January 1846. They took part in the voyage of H.M.S. Herald. The leader of the expedition was the former second in command of the Sulphur, Captain Kellett. The following islands were visited: Gardiner [Gardner], Charles, James, and Chatham [currently Santa María, San Salvador, and San Cristóbal]. What role Berthold Carl Seemann (1825–1872), from England, played on the Galápagos Islands is unclear; he was not mentioned by Wiggins and Porter (1971). His Narrative of the voyage of H.M.S. Herald during the years 1845–1851 … (Seemann, 1853) and The Botany of the voyage of H.M.S. Herald … (Seemann, 1852–1857) give the clear impression that he was the naturalist on the remaining part of the trip, and Goodridge is hardly mentioned. A tragic accident occurred at Atacames in the province of Esmeraldas. Upon boarding the ship after an excursion to areas around Atacames, Seemann wrote, "Several were already in the boat, and I was getting in with the naturalist [Mr. Edmonston] close behind me, when the leg of my trousers lifted the cock of a rifle. The piece went off, sending its charge through the arm of Mr. Whiffin, and making a perfect furrow through the skull of the unfortunate Edmonston. He uttered a slight exclamation, and fell into the water. A man immediately raised him to the surface, but life was gone." Seemann named the genus Edmonstonia after Edmonston and suggested that a new order would probably be needed to accommodate the species—it was that remarkable. Today, E. pacifica is a synonym of Tetrathylacium macrophyllum in the Flacourtiaceae. The expedition left Ecuador for Peru on 28 January 1846, but Seemann returned to Ecuador in 1847. He entered Ecuador overland from Piura via La Peñete, Peru, to Macará, Ecuador. No summary of Seemann's journey in Ecuador has been found, and as both his books are very rare, his itinerary in Ecuador is presented here. For the majority of the place names that Seemann visited he did not use the current spelling. He changed guides frequently, so a particular guide cannot be faulted for the numerous mistakes; they must be attributed to Seemann. On 8 August he traveled from Macará to Soviango [Sabiango] and indicated that he planned to visit Quito. He passed through Sasarango [Zozoranga], Tambo Colosacapi [Colaisaca?], and Cariamango [Cariamanga] on 9 August. Cariamanga was composed of 100 houses, and just north of town was a large cross on the top of a mountain (the cross, or its replacement, is still there). In Gonzanamá he waited a few days to get new guides and mules. He left 16 August for Loja, crossed Río Catamayo to arrive in Loja, or, as he states, previously called Loxa or Zarza in the valley of Cujibamba. He made a trip south of Loja that took him to the Piscobamba valley passing through Vilcabamba. He left Loja on 1 September, crossed the Río Juntas on 2 September, and pushed on to San Lucas, where he observed numerous tree ferns. On 3 September he traveled to Saragura [Saraguro] and on to Oña on 4 September. He left Oña on 5 September and reached Cochopato [Cochapata]. He left Nabón on 7 September and arrived at Cumbi [Cumbe] where he talked to the priest, who remembered William Lobb (1809–1863). [Lobb was a plant collector who had passed through Ecuador a few years earlier. Almost nothing is known about Lobb in Ecuador.] It took Seemann nine days to cover a distance that today is covered in 4–5 hours by automobile. In Cuenca he received news from Captain Kellett. He ordered Seemann to travel immediately to Guayaquil. Seemann did not seem to be in a hurry to leave Cuenca, however. He had time for a party on 12 September and made numerous observations on the city. He left Cuenca on 18 September for Quinoas and passed through Punta de Cajas. On his way down to Tambo Guaicuase and Molleturo, he passed through a thick forest of Podocarpus (remnants of that forest may still be observed today). On 23 September he reached the cave Chacayaque, but due to heavy rain and wind it did not provide much protection. He arrived in Naranjal on 24 September; from Puerto Naranjal he sailed to Guayaquil. He sailed on the Herald from Guayaquil and arrived at Santa Elena on 29 September. On the trip north along the coast he visited Salango on 6 October, Manta on 10 October, Colorado (not to be confused with Santo Domingo de los Colorados) and Monte Cristi on 13 October, Punta Galena [Ballena?] on 16 October, and Río Súa and Esmeraldas on 23 October. The Herald expedition left Ecuador on 25 October when they sailed to Tumaco in Colombia.

Karl Theodor Hartweg (1812–1871), from England, traveled in Ecuador from 1841 to 1842 and collected about 900 plants, most of them with 20 duplicates; the first sets are deposited in London (K and BM). Hartweg worked on behalf of the Horticultural Society of London, of which George Bentham was the president (McVaugh, 1970). He was to collect seeds of plants that would grow in England, but he was allowed to collect herbarium specimens and sell the collections via the society. Bentham described all new species discovered by Hartweg in Plantae Hartwegianae (1839–1857). McVaugh (1970) provided a detailed introduction to the work in the facsimile edition. His introduction contains a complete itinerary of Hartweg's travel and dates of publication for the fascicles. In brief, Hartweg landed in Guayaquil on 13 March 1841, traveled by sea to Santa Rosa and from there inland to Loja, arriving 20 June. He spent more than four months in Loja and surrounding areas and left for Cuenca by 7 November 1841. He stayed in Cuenca until 22 January 1842; however, almost all his plants from Cuenca were lost. Riobamba was his next headquarters, where he stayed until 17 March 1842. He indicated that the trip to Quito followed a "very uninteresting road." The same road has been called the avenue of volcanoes and is among the most scenic roads in the Andes. Hartweg's impression indicated either that the area was already completely converted into farmland so that he did not find anything to collect, or that he traveled the distance during days with low cloud cover. Hartweg stayed in Quito four months and left for Popayán, Colombia, on 21 July 1842.

Gustav Karl Wilhelm Herman Karsten (1817–1908), from Germany, spent 1844–1856 in South America, mainly in Colombia and Venezuela, but he also made explorations in Ecuador. He collected in Pichincha and also reached Sangay (Acosta-Solís, 1969b). Röhl (1944) mentions the following place names in Ecuador: Ibarra, Quito, Cotopaxi, Hambato [Ambato], and Riobamba. He also seems to indicate that he reached the plains of Putumayo. His collections are deposited at Vienna (W) and Berlin (B).

Joseph Warscewicz Ritter von Rawicz (1812–1866), from Lithuania, gardener and independent plant collector, collected in South America from 1845 to 1853. He specialized in collecting live orchids. The localities he visited in Ecuador, probably in 1851, were: Santa Elena, Jipijapa, Guayaquil, Cuenca, and Loja. His collections were deposited at B, later mostly destroyed, with duplicates at BR, CGE, G, K, KRA, U, and W.

Nils Johan Andersson (1821–1880), from Sweden, was on Galápagos during 11–22 May 1852 and collected 325 numbers. Andersson was the botanist on the frigate Eugenie's voyage around the world under the command of Captain C. A. Virgin. Andersson (1854) published his finds from Galápagos in Om Galápagos-öarnes vegetation.

Joseph Pitty Couthouy (1808–1864), malacologist, from the U.S., collected near Quito in 1855. His collections are at G, GH, F, and NY. He took part in a treasure hunt for the ship San Pedro in the Bay of Cumaná from 1854 and enrolled in the U.S. Navy in 1861 (Dall, 1888). Dall (1888) does not mention anything about his collection of plants, but Barnhart (1965) indicated that he did collect in Ecuador and that many of his collections are at the "Torrey Herbarium," i.e., NY.

Jules Ezechiel Rémy (1826–1893), from France, collected on Pichincha, Cotopaxi, and Chimborazo in November 1856. He scaled Pichincha and made an attempt to scale Chimborazo together with Brenchley. His collections are deposited at P, PC, GH, and maybe at M.

Richard Spruce (1817–1893), from England, spent almost seven years in Ecuador from 1857 to 1863. Spruce was primarily a bryologist, but collected numerous higher plants. His diary and letters, edited and condensed by A. F. Wallace, and published as Notes of a Botanist on the Amazon and Andes (1908), offer one of the most fascinating accounts of the traveling conditions of the time. Guzmán's map of the Llangantes as well as a map of Spruce's travels are reproduced therein. His collections are deposited primarily in London (K and BM), but duplicates are found in many herbaria. Acosta-Solís (1969b) provided a condensed summary of the many localities he visited in Ecuador. Spruce arrived in Ecuador by ascending the Pastaza River in April 1857. He collected mainly in the highlands of the provinces of Tungurahua, Chimborazo, León (now Cotopaxi), and Pichincha until June 1860. After that he collected in Bolívar, Los Ríos, and, particularly, around Guayaquil until the end of 1863 when he left for Peru.

Moritz Friedrich Wagner (1813–1887), a German, botanist and zoologist, collected in North and Central America in 1853–1854 and in Panama and Ecuador in 1857–1860. More specifically, he collected in Ecuador from September 1858 to April 1859. He collected 617 numbers, which are deposited at M, with duplicates at BR and FI. Furthers details of his itinerary can be found in Diels (1937).

Juan Isern y Batlló (1825–1866), from Spain, was a botanist on the Royal Spanish Pacific expedition from 1862 to 1865. The expedition, led by Luis H. Pinzón, included scientists of many disciplines. Miller (1968) described Isern as the most diligent member of the expedition. Isern made approximately 500 collections in the lowlands of Ecuador; no complete estimate is known for the highlands, but he made some 160 collections in Pichincha alone (Miller, 1968). The total number of collections for the expedition was 8,000. His collections are deposited in Madrid (MA), with many duplicates at Göteborg (GB). Cuatrecasas (1935a) provides a brief review of the dates and places visited by Isern in Ecuador. A more complete itinerary can be obtained from Barreiro (1926) or Jiménez (1928). Isern and his companions, following the example of La Condamine, left Ecuador through the Amazonian lowlands and traveled down the Amazon to Manaos, Belém, and Pernambuco where they caught ship to Spain. The trip down to the Amazonian lowland was a major undertaking; they estimated that they needed 200 carriers to transport all their collections and equipment. As a result, their collections were shipped in small quantities as carriers became available. Isern caught an infection during the last part of the expedition and died shortly after returning to Madrid (Miller, 1968).

Gustav Wallis (1830–1878), from Germany, arrived in Ecuador and collected in Chimborazo and Pichincha in 1865. He was in Loja from June 1866 until 1868 when he left for Europe. He returned to Ecuador in 1877. He stayed in Guayaquil, but died soon after a trip to Cuenca in 1878. Wallis was a professional collector employed, at least in part, by J. Linden during the first collecting trip and later by J. Veitch & Sons Ltd. for the second trip. Wallis published a few new species, presumably based on material collected while working for Linden, which resulted in a very personal two-page attack from Linden (1875), who accused Wallis of being a poor and corrupt collector with little knowledge of plants. Wallis's (1875) reply was 18 pages long and took the form of several citations from Linden's letters praising him for his work and the excellent collections.

Another collector of major importance to the understanding of the Ecuadorian flora is the Italian clergyman Luigi Sodiro (1836–1909), who stayed in Ecuador from 1870 to his death. Sodiro used both the Spanish version of his first name "Luis" and the latinized form "Aloisius" in his publications. He taught botany at the Escuela Politécnica and at the Central University in Quito, and in that sense replaced Jameson. Sodiro collected in all parts of the country, with special emphasis on the areas around Quito. Some uncertainty still remains about his collections. His private herbarium contains approximately 20,000 collections (including duplicates) and is deposited at "Biblioteca Aurelio Espinosa Pólit" (QPLS) in Quito. It is accessible for scientific study by appointment. Another set was deposited at the Central University (Q), but it is not clear how complete that set was or is. A considerable part of these collections was sold to the Darwinion Institute (SI) in Argentina. Sodiro distributed large amounts of his collections to herbaria in Europe, particularly Berlin, where many were destroyed during the Second World War. Specimens have, however, turned up in BP, S, and FI, and minor portions are in many other herbaria. Sodiro published treatments on Piperaceae, Araceae, Poaceae, Passifloraceae, and various fern families (Sodiro, A., 1900, 1905a, 1905b, 1906, 1908; Sodiro, L., 1893, 1901, 1902–1903, 1903, 1905a, 1905b, 1906, 1908a, 1908b, 1930). His collections are very important because they often contain type material, but they present a problem to researchers due to the relative inaccessibility of the material. To further complicate the situation, Sodiro numbered the species he collected, not his collections. All collections he made of what he believed to be the same species carry the same number. He changed this system at least once, which makes the logic even more difficult to follow, and confusion may occur between the species numbers and collecting dates.

Alphons Stübel (1835–1904), a German, studied volcanoes in Colombia and Ecuador (Stübel, 1897). The time spent in Ecuador was from April 1870 to October 1874. He made numerous collections, and his itinerary is well established by Diels (1937). He collected mainly in the Andes, and Hieronymus (1895b) described his new species. Most of his collections were lost in Berlin (B).

Benedikt Roezl (1824–1885), a Czech who collected in Mexico and South America. He visited Ecuador only once in 1873, making a single excursion from Guayaquil to Chimborazo. His nephews Eduardo (dates unknown) and Franz Klaboch (?–1879), both Czechs, arrived in Ecuador in 1875 and collected orchids in the provinces of Guayas and Azuay until September 1876. Later, they again visited Ecuador, this time accompanied by their cousin B. Hauda.

Édouard François André (1840–1911), a Frenchman, collected in Ecuador from June to August 1876. He worked in the northern part of the country, apparently occasionally together with Sodiro. Smith (1965) wrote, "André was commissioned by the French government to make a general scientific survey of the Northern Andes. Since André was not reimbursed for the expenses of the expedition, he was forced to recoup by selling his specimens to the Royal Botanical Garden, Kew where they may now be consulted." Smith (1965) detailed his itinerary. In brief, he passed through Tulcán, Quito, Riobamba, and Guayaquil. Smith (1965) also questioned whether or not André collected in Loja. It is quite possible that André never collected south of Riobamba and Guayaquil, despite several collections attributed to him from the province of Loja. We have, for instance, recorded in TROPICOS that André collected at El Cisne in the province of Loja, where he collected Streptosolen jamesonii, a plant endemic to the southern part of the country which is currently widely cultivated as an ornamental. The explanation for this contradiction may be that André hired Hugo A.-C. Poortman (also written Poortmann or Portman), a Belgian, to collect in Loja and Zamora-Chinchipe. All collections from Loja attributed to André may be by Poortman. Many of Poortman's collections are deposited at P, and he made at least 529 collections between 3 March and 9 November 1881. Renner (1993) indicated the dates to be from January to June 1882. Poortman's labels were often handwritten on pre-printed André labels, which is the reason for the confusion. Smith indicated that André returned to Ecuador in 1880, but that he had no knowledge of his itinerary. It is possible that André hired Poortman at that time and that was his sole purpose for visiting Ecuador.

Karl [Carl] Friedrich Lehmann (1850–1903), from Germany, was the German consul in Popayán, Colombia. He made frequent collecting trips to Ecuador from 1876 to 1881, going as far south as the provinces of Loja and El Oro. Diels (1937) gave a complete itinerary for his trips to Ecuador. Lehmann drowned crossing Río Timbique, and it is unknown whether he was murdered or an accident occurred (Anonymous, 1904). His collections were deposited in Berlin (B); there are duplicates in many other herbaria.

Edward Whymper (1840–1911), an English geographer and alpinist, primarily was in Ecuador to scale volcanoes in 1879–1880 (Whymper, 1896; Bonney, 1911). His botanical collections were deposited at BM.

Niels Gustaf Lagerheim (1860–1926), from Sweden, lived three years in Quito, from 1889 to 1892, where he was appointed Professor of Cryptogamic Botany and Director of the Botanical Garden (Barnhart, 1965). It is unclear whether he completely replaced Sodiro. They did not publish or collect together, so perhaps Sodiro retired from teaching around 1889, after having taught for nearly 20 years. Lagerheim published six papers soon after leaving Ecuador (Lagerheim, 1891, 1892, 1893a, 1893b, 1894, 1895) that dealt with botany of vascular plants in Ecuador.

August Rimbach (1862–1943), a German, lived in Ecuador from 1890 until his death, with few trips outside Ecuador. He made at least 832 collections and possibly more than 1,000. Rimbach seems to have made exsiccatae and started two series, one in 1906 and the second in 1932. At least the numbers 1–200 may have been used in both series. He was professor of botany and zoology in Cuenca from 1890 to 1894. From 1894 to 1895, he lived in Guayaquil. Between 1895 and 1900, he was in Germany and the U.S. (Nebraska and California). In 1900, he moved back to Guayaquil where he stayed until 1908. From 1908 to 1910, he lived in Riobamba. Then he went to Montevideo for nine years, but returned to Guayaquil in 1919. From 1921 on, he lived in Riobamba, and his collections from that area and time period are famous for their quality. The specimens are often accompanied by colored pencil drawings of extreme precision and detail. His collections were widely distributed; the first set apparently went to B. His brother Carl Rimbach (1864–1933), a geologist, died in Riobamba as well.

Henrik [Heinrich] Franz Alexander Baron von Eggers (1844–1903), from Denmark, was a professional soldier and botanist. From 1869 to 1885 he enlisted in the Danish army and was stationed on the Virgin Islands. This is where his interest in botany started and where he may have collected his first 14,000 collections; these are deposited in Copenhagen (C). He corresponded extensively with Eugen Warming, but a joint trip to the West Indies resulted in a tragic break between them, which explains why Eggers's later collections from Ecuador went to B; duplicates may be found at K, M, O, and US. He stayed in Ecuador from 1891 to 1897 and made 1,700 collections, particularly from the coastal plain. These were apparently the first collections from Ecuador actually to be numbered in the field, but they were not always given exact localities. Other collectors of this era, with the possible exception of Rimbach, never used a numbering system or their plants were numbered upon arrival at the institution, as in the case of the collections of Lehmann, Hartweg, and Spruce. Numbers from 14,000 to 14,700 were collected in the area of Guayaquil, Puná, and especially Balao. The numbers 14,701–15,700 were collected at El Recreo and Zapotal. Eggers published a single paper on the vegetation of coastal Ecuador (Eggers, 1894). El Recreo was Eggers's hacienda. Hugh Iltis relocated the hacienda in 1977; he found the building still standing, about 10 km north of San Vicente in Manabí province (pers. comm. 1997). Iltis interviewed an elderly woman, who remembered the carriages arriving for the parties given by Baron von Eggers.

A number of collectors visited the Galápagos islands in the latter part of the 18^th century. I will only give very brief information here and refer readers to Wiggins and Porter (1971) for further details. A. Habel (dates unknown), from Austria, collected on Galápagos on 22 July 1868, and in January 1869. A total of 69 numbers were collected. Thomas Hill (dates and nationality unknown) and Franz Steindacher (1834–1919), an Austrian ichthyologist, were on Galápagos from 10 to 19 June 1872, and collected 96 numbers. Steindacher's collections are at W. Franz Theodor Wolf (1841–1921), a German, was on Galápagos in August 1875, and only nine collections remain, as most of his plants were lost in storage in Guayaquil. Wolf was professor of geology and mineralogy at the Central University in Quito from 1870, and he was honored with the title of state geologist in 1875. No botanical collections by Wolf are known from the mainland. Wolf returned to Germany in 1891. Gaetano Chierchia and Cesare Marcacci (dates unknown), from Italy, collected on Galápagos from 21 to 31 March 1884; a total of 44 numbers were collected. Leslie A. Lee (1852–1908), from the U.S., was on Galápagos 5–11 April 1888, where he collected a total of 42 numbers. R. E. Snodgrass (dates and nationality unknown) and Edmund Heller (1875–1939), from the U.S., were on Galápagos from 10 December 1898 until mid-June 1899, where they made 949 collections. Alexander Emmanuel Rodolphe Agassiz (1835–1910), Swiss, spent 28 March to 4 April 1891 on Galápagos and collected 41 numbers. George Baur (dates and nationality unknown), spent three months, 6 June to 6 September 1891 on Galápagos and made 385 collections.

Luis Mille (dates unknown, died in the 1940s), Belgian and a pupil of Sodiro, worked in Ecuador from 1891 until his death. He taught botany in Quito and Riobamba. He collected mainly in the Andes until the 1920s when, due to bad health, he moved to the coastal plain and collected in Guayas and Manabí. Eight hundred ninety collections are preserved in Quito (QCA) and constitute one of the oldest collections deposited in Ecuador in a fully operational herbarium (Jørgensen et al., 1992).

Luis Cordero (1833–1912), an Ecuadorian, wrote Enumeración Botánica de las Principales plantas… del Azuay y de Cañar … (1911). It is uncertain whether Cordero made collections, but if so, none are apparently preserved (see Acosta-Solís, 1969b, for further details).

The 20^th century — The number of collectors in this century surpasses 900, and it is impractical to give detailed accounts of so many people. It is conservatively estimated that the number of collections made prior to 1900 is 50,000, and more than 500,000 collections have been made in the 20^th century. The total number of collections made in Ecuador may be as high as 600,000. The number of collections currently available for study is obviously smaller, considering the destruction of herbaria. These figures have been obtained by estimating the number of collections for each of the 113 most important collectors (Table 2) and an additional 176 collectors. The estimated number of collections for the 289 collectors was 495,000. The total estimate is considerably higher than the 250,000 Harling (1986) estimated, even taking into account collections made since then. From the growth rate of the QCA herbarium in the late 1980s it was estimated that about 15,000 collections were made in Ecuador yearly (Jørgensen et al., 1992). These numbers allow us to look at the number of collections made per year in more detail (Figure 2A). The average number of collections made between 1735 and 1800 was about 17 collections/year on the average. In the period 1801–1860 it rose to 290 collections/year, and in 1861–1930 it grew again, this time to 696 collections/year. Between 1931 and 1975, a significant increase occurred, bringing the total to 3,187 collections/year. Since 1975 the rate of collecting in Ecuador has again increased, to 15,615 collections/year on the average. An increase from about 18 collections per 100 km² at the start of this century to more than 200 collections per 100 km² clearly indicates that a considerable effort has been made to document the botanical diversity in Ecuador. This number surpasses the minimum threshold value for making botanical inventories of 100 collections per 100 km² suggested by Campbell (1989), and Ecuador must be among the best-collected areas in South America. A glance at the map of all georeferenced collections in TROPICOS, on the right front endpaper, confirms that Ecuador is well collected. It also makes clear that collecting is not evenly or randomly distributed, but to a large degree follows such access routes as roads and rivers. The result is that some areas, mainly those close to Quito, field stations, or along roads, are very well collected, while other areas are still almost unexplored.

It would be logical if the rate of publication of species were correlated with that of exploration and collecting. To document that, I have looked at the time of publication of the species recognized in the Catalogue (Figure 2B). The average number of accepted species published per year for the five time periods identified above were: 26 species/year (1753–1800), 58 (1801–1860), 61 (1861–1930), 67 (1931–1974), and 165 (1975–1998). Since 1975, on average, every 95 collections yielded one new species described, and every two days a new species has been described from Ecuador. The average number of endemic species published in the five periods were: 0.1, 4, 12, 17, and 91, respectively. It is still relatively easy to find new species in Ecuador, but the number of collections needed is increasing. Between 1801 and 1860 it took on the average 73 collections to find a new endemic species, but only five collections to find a new species. At present it takes 170 collections, on the average, to find a new endemic species in Ecuador. This suggests that about one in two new species decribed from Ecuador is endemic to the country—known nowhere else. Every fourth day an endemic species is being described from Ecuador. It is clear that numerous species described as endemic do not remain endemic forever. Often the species is later found in other countries, or lumping of the species into already established species can occur. Nevertheless, many new finds are still emerging and there are still many new species to be found; the era of discovery and exploration is far from over in the tropics.